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Ellis, E.C. 2015. Ecology in an Anthropogenic Biosphere. Ecological Monographs in press doi.org: 10.1890/14-2274.1 [FORMATTED PREPRINT]
Ecology in an Anthropogenic Biosphere
ERLE C. ELLIS1*
1
Department of Geography & Environmental Systems, University of Maryland, Baltimore County, Baltimore, MD, 21250 USA.
Abstract. Humans, unlike any other multicellular species in Earth’s history, have emerged as a global
force that is transforming the ecology of an entire planet. It is no longer possible to understand, predict, or
successfully manage ecological pattern, process or change without understanding why and how humans
reshape these over the long-term. Here, a general causal theory is presented to explain why human societies
gained the capacity to globally alter the patterns, processes and dynamics of ecology and how these
anthropogenic alterations unfold over time and space as societies themselves change over human
generational time. Building on existing theories of ecosystem engineering, niche construction, inclusive
inheritance, cultural evolution, ultrasociality, and social change, this theory of anthroecological change
holds that sociocultural evolution of subsistence regimes based on ecosystem engineering, social
specialization and nonkin exchange, or “sociocultural niche construction”, is the main cause of both the
long-term upscaling of human societies and their unprecedented transformation of the biosphere. Human
sociocultural niche construction can explain, where classic ecological theory cannot, the sustained
transformative effects of human societies on biogeography, ecological succession, ecosystem processes,
and the ecological patterns and processes of landscapes, biomes and the biosphere. Anthroecology theory
generates empirically testable hypotheses on the forms and trajectories of long-term anthropogenic
ecological change that have significant theoretical and practical implications across the subdisciplines of
ecology and conservation. Though still at an early stage of development, anthroecology theory aligns with
and integrates established theoretical frameworks including social-ecological systems, social metabolism,
countryside biogeography, novel ecosystems and anthromes. The "fluxes of nature" are fast becoming
"cultures of nature". To investigate, understand, and address the ultimate causes of anthropogenic
ecological change, not just the consequences, human sociocultural processes must become as much a part
of ecological theory and practice as biological and geophysical processes are now. Strategies for achieving
this goal and for advancing ecological science and conservation in an increasingly anthropogenic biosphere
are presented.
Key words: Anthropocene; human dominated ecosystems; human impact; disturbance; ecological
inheritance; extended evolutionary synthesis; anthropogenic landscapes; biodiversity; anthrosequence;
anthrobiogeography; human ecology; archaeology; natural history.
“it would be difficult, not to say impossible, to draw a natural line between the activities of the human tribes which
presumably fitted into and formed parts of "biotic communities" and the destructive human activities of the modern
world.” (Tansley 1935)
“the vast bulk of the impact that human beings have made on this planet has undoubtedly resulted directly from socially
transmitted knowledge.” (Odling-Smee and Laland 2012)
“changes in human societies over the past 12,000 years can be understood as constituting a single complicated earth-wide
event of spiraling globalization.” (Chase-Dunn and Lerro 2013)
INTRODUCTION
Human societies have been altering ecological and
evolutionary processes across the Earth for millennia (Butzer
1982, Redman 1999, Grayson 2001, Kirch 2005, Barnosky 2008,
Ellis 2011, Doughty 2013, Smith and Zeder 2013, Barnosky
2014). As behaviorally modern Homo sapiens spread out of
Africa more than 50,000 years ago (Klein 2013), their advanced
hunter gatherer societies helped to cause the extinction of more
than half of Earth’s mammalian megafauna, yielding trophic
Original manuscript received 26 November 2014; accepted with
revisions 2 February 2015; accepted 12 March 2015. Corresponding
Editor: Aaron Ellison
*
Present address: Department of Geography & Environmental Systems,
University of Maryland, Baltimore County, 1000 Hilltop Circle,
Baltimore, MD 21250 USA. E-mail: [email protected]
cascading effects on ecosystems coupled with the direct effects
of landscape burning to enhance hunting and foraging success
(Grayson 2001, Barnosky 2008, Estes et al. 2011, Doughty 2013,
Barnosky 2014). More than 10,000 years ago, agricultural
societies accelerated these early defaunation and land clearing
processes, ultimately replacing them with even more novel
ecological transformations, including the culture of
domesticated species, widespread soil tillage, sustained societal
growth, and ever increasing scales of material exchange, leading
to globally significant transformation of the terrestrial biosphere
by at least 3000 years before the present time (FIG 1A; Kirch
2005, Ellis 2011, Ellis et al. 2013b, Smith and Zeder 2013).
Human societies have now caused global changes in
atmospheric composition and climate (IPCC 2013), hydrology
(Vörösmarty and Sahagian 2000), geomorphology (Wilkinson
2005, Syvitski and Kettner 2011), fire regimes (Bowman et al.
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Ellis, E.C. 2015. Ecology in an Anthropogenic Biosphere. Ecological Monographs in press doi.org: 10.1890/14-2274.1 [FORMATTED PREPRINT]
FIG. 1. Human transformation of the terrestrial biosphere. (A) Time period of first significant human use of land for agriculture and settlements in
regions with sustained use (years BP), urban and densely settled areas in year 2000, and percent recovery from peak land use in regions without
sustained use (modified from Ellis et al. 2013b). (B) Anthropogenic biomes (anthromes; year 2000; modified from Ellis et al. 2010). Anthromes are
organized into five levels in the legend. Eckert IV equal area projection.
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Ellis, E.C. 2015. Ecology in an Anthropogenic Biosphere. Ecological Monographs in press doi.org: 10.1890/14-2274.1 [FORMATTED PREPRINT]
2011), and other Earth systems (Zalasiewicz et al. 2012). Human
societies have caused widespread species extinctions (Barnosky
2008, Dirzo et al. 2014, Pimm et al. 2014) and species invasions
(Vitousek et al. 1997a, Ricciardi 2007, Lockwood et al. 2013),
changes in the local and global patterns of net primary
production (Vitousek et al. 1986, Krausmann et al. 2013) and in
the local and global biogeochemical cycles of carbon, nitrogen,
phosphorus and other elements (Vitousek et al. 1997b,
Falkowski et al. 2000, Galloway et al. 2004, Elser et al. 2007).
Humans have also introduced a wide array of entirely new
ecological processes to the Earth system (Ellis and Haff 2009)
including species domestication by artificial selection (Larson et
al. 2014), direct genetic modification of organisms (Dale et al.
2002), large-scale combustion of fossilized photosynthates
(Boden et al. 2012), artificial lighting (Longcore and Rich 2004),
and the chemical synthesis of reactive nitrogen (Gruber and
Galloway 2008) together with a vast number of artificial
chemicals, plastics and other synthetic materials, many of which
are used to control other species (Alloway and Ayres 1997,
Corcoran et al. 2014). Humans have facilitated the release and
utilization of nuclear energy and radionuclides (Harrison et al.
2011, Hancock et al. 2014, Zalasiewicz et al. 2015). Humans
engage in annual soil tillage and produce large-scale earthworks,
channels, tunnels, and boreholes (Richter et al. 2011, Edgeworth
2014, Zalasiewicz et al. 2014) including massive vertical built
structures (Frolking et al. 2013), and artificial structures
interconnected across continents (e.g. canals, roads, railways,
(Forman and Alexander 1998, Verburg et al. 2011). Humans
have also introduced increasingly rapid, massive and
mechanized aerial, terrestrial and hydraulic transport of material
and biota across the Earth (Mooney and Cleland 2001,
Steinberger et al. 2010, D'Odorico et al. 2014) and these
transport processes are beginning to extend beyond Earth’s
atmosphere (Crowther 2002, Gorman 2014). The rates and
scales of these processes continue to accelerate driven by
increasing use of non-biological energy and the mechanization
and automation of human activities, from food production and
construction to trade and communications (Smil 2008, Baccini
and Brunner 2012, Brynjolfsson and McAfee 2014).
Taken together, these anthropogenic global environmental
changes have been characterized as the emergence of humans as
a “great force of nature” that is transforming the Earth system,
shifting the planet into a new epoch of geologic time; the
Anthropocene (Crutzen 2002, Steffen et al. 2007, Ellis 2011,
Steffen et al. 2011, Zalasiewicz et al. 2012, Smith and Zeder
2013, Barnosky 2014). Whether or not the Anthropocene is
formally recognized, there is no question that the scale, rate,
intensity and diversity of anthropogenic environmental changes
are unprecedented in comparison with those caused by any prior
multicellular species. The question for ecology is not whether,
when, or even how humans have transformed the biosphere but
rather, why?
Evolutionary theorists and social scientists have made
substantial progress towards explaining the exceptional growth
and development of human societies and their unprecedented
capacity for environmental transformation, especially in
archaeology, anthropology and sociology (e.g. Butzer 1982,
Laland et al. 2000, Kirch 2005, Nolan and Lenski 2010, ChaseDunn and Lerro 2013). Yet ecology remains without a widely
accepted causal theory that can explain how a single
multicellular species gained the capacity to transform an entire
planet– though such theories have a long history (e.g. Marsh
1865, de Chardin 1955, Vernadsky 1998) and are of increasing
interest to ecologists (e.g. Barnosky 2008, Collins et al. 2011,
Steffen et al. 2011, Barnosky et al. 2012, Costanza et al. 2012,
Smith and Zeder 2013, Malhi 2014).
It is no longer possible to explain or predict ecological
patterns or processes across the Earth without considering the
human role in these (Ellis and Ramankutty 2008, Ellis and Haff
2009, Barnosky et al. 2012). More than three quarters of the
terrestrial biosphere has already been transformed into
anthropogenic biomes (anthromes) by human populations and
their use of land (FIG 1B; Ellis and Ramankutty 2008). In a
biosphere increasingly transformed by human societies, ecology
cannot advance as a predictive science without gaining the basic
theoretical tools needed to investigate and understand the
ultimate causes, not just the consequences, of human
transformation of ecological pattern, process and change.
TOWARDS A GENERAL THEORY OF
ANTHROPOGENIC ECOLOGICAL CHANGE
This paper introduces a general causal theory of long-term
anthropogenic ecological change (“anthroecology theory”)
based on a synthesis of contemporary evidence and theory
across the ecological, evolutionary, and social sciences. The
evidence will show that the ultimate causes of unprecedented
human transformation of the biosphere are social and cultural;
not biological, chemical or physical. As a result, an
understanding of human sociocultural processes is central to
understanding anthropogenic ecological change. Disciplinary
barriers to this understanding are formidable (Laland and Brown
2011). Yet there is no other way forward.
Human societies have emerged as a global force that is
reshaping the biosphere, atmosphere, hydrosphere and
lithosphere (Steffen et al. 2007). To understand this massive and
sustained human transformation of Earth’s ecology it is
necessary to consider human societies as a global force capable
of interacting with and reshaping ecology across the Earth in
ways analogous to that of the climate system (Ellis and Haff
2009, Lucht 2010, Steffen et al. 2011). Just as a causal
understanding of climate and weather is needed to understand
the ecological patterns, processes and dynamics produced by
these across the Earth, a causal understanding of human
sociocultural processes- analogous to a “human climate and
weather”- is required to understand the long-term ecological
patterns and processes resulting from sustained interactions with
human societies.
Social-ecological systems (SES) theory has developed into a
useful framework for understanding, predicting and addressing
the dynamics of a "human weather" in which human societies
interact directly with populations and ecosystems locally and
regionally with direct feedbacks over years to decades (Folke et
al. 2005, Alessa and Chapin 2008, Carpenter et al. 2009, Chapin
III et al. 2011, Collins et al. 2011, Levin et al. 2013). SES
frameworks have become especially useful for understanding
the dynamic interplay of coupled social and ecological systems
in practical settings generally with the aim of promoting resilient
system interactions together with stakeholders (Folke 2006,
Chapin III et al. 2011, Collins et al. 2011)
Here the goal is more basic: to explain the emergence and
long-term dynamics of the “human climate system” that has
been reshaping the terrestrial biosphere for more than 50,000
years and that will likely continue reshaping it into the
foreseeable future. This explanation has two parts. The first is to
explain how human societies initially gained their
unprecedented capacity to transform ecological and evolutionary
processes. The second is to explain how this capacity has scaled
up and changed as a transformative force on ecology as human
societies themselves have changed and diversified over human
generational time, from small bands of hunter gatherers to
globalized industrial societies. To gain this understanding it is
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Ellis, E.C. 2015. Ecology in an Anthropogenic Biosphere. Ecological Monographs in press doi.org: 10.1890/14-2274.1 [FORMATTED PREPRINT]
TABLE 1. Definition of terms, in order of appearance in the text. Terms in italics are either newly defined or redefined for use here.
Term
Definition
Altered or influenced by human sociocultural systems. Anthropogenic alteration of ecosystems and evolutionary processes may be
Anthropogenic direct or indirect, intended or unintended, and have neutral, beneficial, detrimental or combined consequences for humans and/or
non-humans.
Human populations engaging in a suite of complex and symbolic social behaviors that emerged fairly rapidly approximately 60,000
Behaviorally modern years ago in Africa as evidenced by major shifts in tool sophistication, diversity and standardization, personal ornaments, pigments,
humans symbolic inscriptions, a broadening of the subsistence base, increasing settlement and population size and long-distance trade.
Linkage with anatomical or other genetic adaptations remains controversial (Nowell 2010).
Anthrome, Globally significant ecological patterns produced by sustained direct human interactions with ecosystems (Ellis and Ramankutty
anthropogenic biome 2008). A globally significant anthroecosystem pattern.
Anthropogenic The biosphere as reshaped by sustained direct interactions with humans (Ellis and Ramankutty 2008). The global anthroecosystem.
biosphere The system formed by the interactions of a world system and the biosphere.
Sociocultural
Processes sustaining sociocultural systems, including both internal processes and those connected with the natural world.
processes
A system composed of a human population and its social institutions, culture, and material products: a human society (Nolan and
Sociocultural system, Lenski 2010) or human system. Sociocultural systems generate, transmit, reproduce, select for, and accumulate cultural, material
human system and ecological inheritance. Sociocultural systems can be identified at multiple scales, from small bands and tribes, to small social
groups within a society to an entire world system.
Natural processes,
natural systems, Processes, systems or states unaltered by human sociocultural systems, e.g. natural ecosystems or natural landscapes; “pristine”.
natural state
Organismal alteration of ecological patterns and processes in ways that confer heritable advantages and/or disadvantages to
individuals or populations (ecological inheritance). Niche construction likely applies to all species in some form. Ecosystem
Niche construction
engineers engage in niche construction only to the extent that their environmental alterations yield heritable consequences.
Examples: soil acidification by nitrifying bacteria, termite nest building, beaver dams, rice paddy systems, air pollution.
Heritable ecological patterns and processes capable of conferring adaptive advantages and/or disadvantages to individuals,
populations and sociocultural systems. The constructed niche that is inherited. Examples: degraded or improved habitats, nests,
Ecological inheritance
dams, burrows, buildup of nutrients or toxins in soils or water, eroded soils, co-evolved species (flowers of pollinators),
domesticates, planted forests.
Extended
An extension of the Modern Synthesis of evolutionary theory to include non-genetic inheritance together with genetic inheritance
Evolutionary
(“inclusive inheritance”). Also called the “Inclusive Evolutionary Synthesis (Danchin et al. 2011, Danchin 2013).
Synthesis (EES)
Non-genetic “The part of variation in a trait that is transmitted to offspring through mechanisms other than genetic variation.” (Danchin 2013).
inheritance Includes epigenetic, parental, cultural and ecological characteristics transmitted across generations as part of the EES.
The degree to which phenotypic characteristics are transmitted between generations, whatever the mechanism of transmission.
Includes both genetic and non-genetic characteristics inherited across generations, and processes of inheritance from parent to
Inclusive heritability
offspring (vertical inheritance), from older to younger generations (oblique transmission) as defined in the EES (Danchin et al.
2011)
The inheritance of both genetic and non-genetic phenotypic information between generations, whatever the mechanism of
Inclusive inheritance
transmission. Phenotypic inheritance as defined in the EES (Danchin et al. 2011)
Information transmitted across generations through social learning (Danchin et al. 2011). Examples: bird song, tool-use,
Culture subsistence strategies, languages, social roles and institutions, advanced tool-making “recipes” and other complex technologies
(Mesoudi and O’brien 2008).
Cultural information enabling or enhancing the behavioral capacity of organisms beyond their biological capabilities. Technology
enables the production and use of tools and machines, but is not the same as the tools and machines themselves, which are defined
Technology
as material culture. Examples: tool use, tool making recipes, control of fire, techniques for machine and computer manufacture,
computer software.
Information transmitted across generations through social learning by vertical or oblique transmission processes. Examples:
Cultural inheritance
subsistence strategies, tool-making knowledge, languages, family planning.
Processes by which variations in cultural information are produced, transmitted, selected for, change in frequency and accumulate
across generations. Cultural macro-evolution explains variations across societies; cultural micro-evolution explains variations
Cultural evolution within societies and social groups. This definition is consistent with the EES and other theory explaining cross-generational cultural
inheritance, and is not the definition used in theory explaining horizontal transmission of cultural information within generations,
such as memetics.
Cultural niche Alteration of ecological patterns and processes by organisms through socially learned behaviors that produce heritable advantages
construction and/or disadvantages to individuals or populations (ecological inheritance). Examples: tool-use, agriculture, pollution.
Dependence on nonkin social cooperation for survival and the biological and sociocultural adaptations enabling this. e.g. (Hill et al.
Ultrasociality
2009, Tomasello 2014).
Processes of ecosystem and material culture engineering that require non-kin social cooperation to enact, including collaborative
Cooperative
manufacture of tools and other material culture, large scale landscape modifications for hunting, irrigation, and transportation
engineering
infrastructure, and complex agricultural and industrial systems.
Regime shifts in the scale, in terms of number of interacting individuals sustained within a sociocultural system, from smaller to
Social upscaling
larger. Transitions from smaller scale societies (e.g. hunter gatherers to horticultural societies; Table 5)
Substitution of one form of energy for another, such as the substitution of fire or animal traction for human labor in clearing land,
Energy substitution
or use of nuclear energy to substitute for coal.
The heritable sociocultural, material and ecological conditions within which human individuals, groups and populations reproduce
Sociocultural niche
and sustain themselves. Applies only to species dependent on socially learned exchange relations.
Alteration of sociocultural, ecological or material patterns and processes by human individuals, groups or populations through
Sociocultural niche socially learned behaviors, exchange relations and cooperative engineering in ways that confer heritable benefits and/or detriments
construction (SNC) to these individuals, groups or populations. Examples: agriculture, pollution, the marketplace, electrical grids, nuclear power,
genetic engineering.
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TABLE 1. Continued.
Term
Definition
Shared intentionality The ability to interpret the intentions of others and to act cooperatively with these intentions, a capacity for intentionality
and social cognition not shared by any other species (Tomasello et al. 2005).
Sociocultural Processes by which variations in cultural information are produced, transmitted, selected for, change in frequency and
evolution accumulate across generations and across sociocultural systems. Sociocultural evolution includes processes of change not
only in cultural information relevant to individuals, but also the cultural information, organization and exchange relations
that structure societies themselves. As with biological evolution, cross-generational changes in the frequencies of culturally
based individual and population behaviors within and across sociocultural systems are influenced by their heritable
advantages and/or disadvantages to individuals, populations and groups.
Material culture Artificial materials produced by sociocultural systems, ranging from stone tools and other small artifacts to the advanced
technologies and built infrastructure of industrial societies; these are key evidence in archaeology (Eerkens and Lipo
2007). Examples: tools, artificial materials and chemicals, built structures, roads and other infrastructure.
Anthroecosystem, A system composed of sustained interactions among human sociocultural systems, non-human biota and abiotic
anthropogenic environment (Fig. 2). A sociocultural system interacting with an ecosystem.
ecosystem
Material inheritance Heritable artificial material products of behaviorally modern human societies that confer advantages or disadvantages to
human individuals, populations, or sociocultural systems and that cannot be produced outside of human societies.
Examples: tools, machines, artificial materials and chemicals, built structures, roads and other infrastructure. Similar to
“material culture”, but only when heritable and selectable.
Subsistence strategy Behaviors that sustain organisms, including foraging strategies, ecosystem engineering, and subsistence exchange.
Subsistence exchange Exchange of materials, labor, energy and/or information among individuals that assist in their survival and reproduction.
Subsistence regime Socially learned subsistence strategies implemented within and by sociocultural systems. Sociocultural subsistence regimes
may be simple or complex (composed of multiple interacting subsistence strategies) and may or may not produce
ecological, cultural and/or material inheritance. Examples: tool-making techniques, cultivation of crops, trading networks,
craft industries (more in Table 6).
World system A sociocultural system formed from interacting sociocultural systems. Analogous to the interacting ecosystems comprising
the biosphere.
Abiotic energy Energy not derived from any biological process, past or present, including solar, wind, geothermal, tidal and nuclear.
Excluding fossil fuels and biological products of any kind.
Land use Intentional alteration of terrestrial ecosystems by sociocultural systems, including harvesting, extracting, cultivating,
grazing, building on, restructuring, managing, conserving and restoring ecosystem pattern and process. Land use produces
ecological inheritance such as the intentional direct benefits of food production by agricultural ecosystem engineering and
detrimental effects from overhunting, soil erosion and pollution.
Wildlands Landscapes or ecosystems that are not currently intentionally altered by sociocultural systems. Free of agriculture and
permanent human settlements. Wildlands anthromes are defined as having no evidence of land use or human populations
(Ellis et al. 2010)
Seminatural lands, Landscapes or ecosystems which show low levels of intentional alteration by sociocultural systems. Seminatural anthromes
seminatural are defined as having <20% land use for agriculture and settlements (Ellis et al. 2010).
ecosystems
Used lands Lands significantly altered intentionally by sociocultural systems. OR, in anthrome classification, landscapes with >20%
land in use for agriculture and settlements (Ellis et al. 2010).
Novel ecosystem An ecosystem altered permanently by interactions with sociocultural systems.
Anthrosequence Anthrosequences depict hypothetical patterns in ecological processes caused by variations in sociocultural niche
construction by different societies acting on a given biome, analogous to the patterning of ecological processes by time
(chronosequence), terrain (toposequences), and climate (climosequences). FIG 5 is an example of a woodland biome
anthrosequence.
Telecoupling The capacity of human populations to exert ecological effects over long distances by means of nonkin exchange relations,
especially the demands for materials, biota and energy expressed through markets (Liu et al. 2013).
Decoupling Increasing the efficiency of resource production or extraction in ways that reduce direct utilization or harvesting of primary
resources and transformation of natural ecosystems (Ausubel and Waggoner 2008, Fischer-Kowalski and Swilling 2011).
Examples, the use of less fossil fuel per mile traveled by a vehicle, the substitution of fossil fuels for biomass energy –
reducing use of land for biomass production, or production of food by more productive agricultural systems instead of
hunting and foraging.
necessary to see beyond the short-term local and regional
dynamics of the “human weather” to focus on the “human
climate system”; the long-term processes by which human
societies act as a global force transforming Earth’s ecology.
A detailed global understanding of social-ecological changes
over the past 50,000 years is certainly beyond the scope of any
single published work. The ultimate purpose here is to guide
ecological science towards a general causal framework
explaining why long-term societal changes reshape ecology and
evolution, with the goal of generating ecologically useful
hypotheses. To build this framework, terms are borrowed from
the social and evolutionary sciences and some are defined
specifically here; Table 1 provides definitions of key terms. A
practical distinction is made here between “natural” patterns and
processes, defined as those unaltered by humans, and
anthropogenic patterns and processes which have been altered,
or introduced de novo by human sociocultural systems,
representing the “human climate system”.
The way forward begins with a simple question. Why and
how did a single species of hominin, so biologically and
ecologically similar to others in its genus (Sterelny 2011, Antón
et al. 2014), originate, scale up (social upscaling; Table 1), and
sustain for generations the culturally complex societies capable
of transforming an entire planet? The answer begins with a
review of recent evolutionary theory explaining how socially
transmitted behavioral strategies that transform environments
and enable societies to scale up through processes of social
specialization and nonkin social exchange can evolve across
generations by enhancing the adaptive fitness of individuals,
progeny, populations, and societies. As will be seen, the
emergence of behaviorally modern human societies as a global
force transforming the biosphere has resulted from an
unprecedented ability for these societies to scale up socially and
to accumulate capacity for ecosystem engineering while
harnessing non-human and even abiotic energy in this process.
To understand why and how a single biological species
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Ellis, E.C. 2015. Ecology in an Anthropogenic Biosphere. Ecological Monographs in press doi.org: 10.1890/14-2274.1 [FORMATTED PREPRINT]
gained the capacity to transform the biosphere requires an
evolutionary theory of anthroecological change based on
“sociocultural niche construction” (Table 1), an integrative
theory constructed from existing theories of ecosystem
engineering, niche construction, ecological inheritance, inclusive
inheritance, cultural inheritance, cultural evolution, and
ultrasociality. Elements of this theory are outlined in Table 2
and presented in detail in the remainder of this section.
Background readings are listed in Table 3.
Ecosystem engineering, niche construction,
and ecological inheritance
Many species transform their physical environments by
processes of “ecosystem engineering” (Jones et al. 1994, Wright
and Jones 2006, Erwin 2008, Jones et al. 2010, Matthews et al.
2014). Yet the evolutionary consequences of this transformation
have only recently been incorporated into ecological theory
(Lewontin 1983, Odling-Smee 1988, Matthews et al. 2014).
Niche construction theory, introduced in the 1990s, links
environmental alterations by organisms to changes in their
adaptive fitness, and that of their progeny, and also that of other
species within these altered environments (Odling-Smee et al.
1996, Laland et al. 1999, Odling-Smee et al. 2003c). Niche
construction theory challenges one of the most basic
assumptions underlying modern evolutionary theory, that
organisms must adapt solely within environmental conditions
that they cannot alter, and replaces this with a “two-way street”
(Ellison 2004) in which organisms alter environments and must
in turn adapt to these altered environments, generating complex
feedbacks in both ecological and evolutionary processes. The
theory has strong empirical and mathematical support (Laland et
al. 1999, Odling-Smee et al. 2003c), applies broadly across taxa,
and is increasingly well connected with existing theory in
ecology and evolution (Odling-Smee et al. 2003b, Erwin 2008,
Odling-Smee et al. 2013, Matthews et al. 2014, Scott-Phillips et
al. 2014).
Niche construction theory holds that by enhancing or
degrading environments in ways that increase or decrease the
adaptive fitness of later generations, or that of other species
interacting with a given environment, organisms produce an
“ecological inheritance” (Table 1; (Odling-Smee et al. 2003c).
Well documented examples of ecological inheritance include the
positive fitness benefits received by a species through the
construction of protective burrows, nests and webs and the
buildup of nutrients and organic material in soils, and also the
negative fitness, or detrimental effects, of depleting soil
nutrients, soil erosion and the buildup of harmful chemicals in
soil and water (Odling-Smee et al. 2003c). Given the many
pathways by which species may alter environments and their
varied consequences, it is useful to first characterize these as
either direct (environmental alteration by a species directly
affects a species- the same or not; e.g. toxic accumulation in
soils) or indirect (environmental alteration by a species affects
the same or another species by altering an additional
environmental process, e.g. production of flammable leaves
increases fire frequencies causing a fire effect), and then to
characterize their effects on the adaptive fitness of the acting
species and/or other species as either beneficial, detrimental, or
compound (beneficial and detrimental) ecological inheritance.
For example, fires produced by the buildup of flammable leaves
would simultaneously yield indirect beneficial ecological
inheritance to a fire-dependent plant species and indirect
detrimental ecological inheritance to a fire intolerant species in
the same environment.
One major evolutionary consequence of ecological
TABLE 2. Elements of human sociocultural niche construction (SNC) theory.
Points in bold correspond to sections in the text. Terms in italics are defined
in Table 1.
1. Ecosystem engineering, niche construction and ecological inheritance.
1.1 Allogenic ecosystem engineers engage in environment altering behaviors,
including nest building, bioturbation and agriculture.
1.2 To the extent that ecosystem engineering behaviors produce adaptive
advantages and disadvantages, this constitutes niche construction, and
produces ecological inheritance.
2. Inclusive inheritance, cultural inheritance and The Extended
Evolutionary Synthesis (EES).
2.1 Natural selection acts on a combined vector of inheritances (inclusive
inheritance) comprising not only genetic and epigenetic inheritance
(heritable alterations of gene expression) but also nongenetic
inheritances, including parental effects, cultural inheritances and
ecological inheritances.
2.2 Inheritances may be transmitted vertically (parent to offspring), obliquely
(older to younger) and horizontally (across individuals). Natural selection
can act on all of these transmission pathways.
2.3 When environments remain stable across generations, genetic and
epigenetic inheritances tend to be the predominant form of adaptive trait
transmission.
2.4 When environments change substantially between generations,
nongenetic inheritances, especially cultural and ecological inheritances,
become increasingly important for adaptive trait transmission.
2.5 Ecosystem engineers cause environmental change. As a result, ecological
and cultural inheritances are favored mechanisms for adaptive trait
transmission in ecosystem engineering species, especially those with
culturally inherited traits for ecosystem engineering, like humans.
2.6 To the extent that socially learned ecosystem engineering behaviors
produce adaptive benefits or detriments, this is cultural niche
construction.
3. Cultural evolution.
3.1 Cultural inheritances evolve by processes of natural selection acting on
traits transmitted vertically, obliquely and horizontally.
3.2 Human cultural inheritances include languages, technologies, social
strategies for ecosystem engineering, foraging, societal organization and
interaction, material and labor exchange, warfare, and defense.
3.3 Cultural traits can evolve far more rapidly than genetic and epigenetic
traits owing to natural selection acting on horizontal inheritances within a
single generation, because selection can act on novel combinations of
preexisting cultural inheritances (e.g. the “recipe” for manufacturing a
tool”), and because selection can also act at the scale of interacting social
groups or societies.
4. Human ultrasociality and the human sociocultural niche (Table 4).
4.1 Behaviorally modern humans are ultrasocial, with unrivalled capacity for
high fidelity transmission of cultural traits, exemplified by human
capacity for language together with unrivaled capacity to form, sustain,
and depend for survival on complex nonkin social relationships and
cooperative material and labor exchanges within and across social groups
and societies.
4.2 Behaviorally modern humans occupy a sociocultural niche, as cultural
traits enable individuals to sustain themselves and their progeny within
social groups and societies (sociocultural systems) and to form and
sustain the structure and functioning of both the sociocultural systems and
the engineered ecosystems, exchange networks and other subsistence
regimes that support human populations,.
4.3 The human niche evolves in response to changes in social organization
brought about both by cultural evolution and the environmental changes
caused by cultural niche construction.
4.4 The unique human capacity for shared intentionality enables individual
choices to scale up to larger group decisions; individual humans and
social groups act as agents deciding and acting within culturally inherited
but dynamic sociocultural systems that emerge from these interactions.
5. The ratchet effect and runaway cultural niche construction. Cultural
traits have overwhelmed genetic traits in shaping the human niche, in part
because of more rapid processes of cultural evolution, including horizontal
trait selection, the ratchet effect, and runaway cultural niche construction.
6. Human sociocultural niche construction (SNC). Long-term changes and
diversification of the human niche and the upscaling of human societies and
their capacity to transform the biosphere (Table 4) can be explained by
combining cultural niche construction, culturally mediated social
organization, and cultural evolution into a single theory of sociocultural
niche construction.
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inheritance is that environment altering traits that produce
fitness benefits for a given species (e.g. burrowing, nest
building, web construction, nutrient-rich leaves or allelopathic
chemicals) or that help a species adapt to the detrimental effects
of environmental changes (e.g. enhanced nutrient uptake or
recycling, metabolism of toxics) or that increase the beneficial
effects of these changes (e.g. burrow and nest re-use, root
systems adapted to nutrient-rich soils), will be selected for in
these altered environments and therefore increase or decrease in
frequency in populations by processes of natural selection,
causing these “recipient traits” to evolve over generational time
(Odling-Smee et al. 2013). The second, and perhaps even
greater consequence of ecological inheritance is that species
engaging in niche construction tend to increasingly alter the
environments and adaptive fitness experienced by other species,
generating broad changes in the ecology and evolution of entire
communities (Odling-Smee and Laland 2012). For example,
niche construction by bioturbation, biostabilization, bioerosion,
and bioconstruction are increasingly recognized as globally
significant forces shaping both biotic and geomorphic processes
across the Earth over the long term (Erwin 2008, Corenblit et al.
2011). The oxygenation of Earth’s atmosphere by
photosynthetic microorganisms is perhaps the ultimate example
of the global transformative power of niche construction
(Odling-Smee et al. 2003c).
The behavioral traits of active allogenic ecosystem engineers,
like dam-building by beavers and nest construction by termites,
with their direct adaptive fitness benefits, are among the clearest
examples of how ecological inheritance can drive the evolution
of environmental altering traits (Matthews et al. 2014). To
understand the evolution of ecosystem engineering by humans,
“the ultimate ecosystem engineers”, the effects of directly
beneficial ecological inheritance are paramount (Laland et al.
2000, Odling-Smee et al. 2003a, Smith 2007b).
Inclusive inheritance, cultural inheritance and the Extended
Evolutionary Synthesis (EES)
Ecological inheritance is now integrated with other forms of
non-genetic inheritance including heritable modifications of
gene activation (epigenetic inheritance), heritable parental
effects, and the inheritance of cultural traits, into an “Extended
Evolutionary Synthesis” (EES; Table 1) that expands the
“Modern Synthesis” beyond genetics to explain the evolution of
complex phenotypic traits across a variety of taxa
(Bonduriansky and Day 2009, Danchin et al. 2011,
Bonduriansky 2012, Danchin 2013, Mesoudi et al. 2013). The
EES combines all forms of heritable phenotypic information into
a vector of “inclusive inheritance” (Table 1) that can be
transmitted across generations from parent to progeny (vertical
transmission), from older to younger generations (oblique
transmission), and among siblings, kin and nonkin within a
generation (horizontal transmission) (Danchin et al. 2011,
Danchin 2013). In this way, evolution by natural selection has
been extended beyond the vertical genetic inheritance of higher
organisms to the oblique and horizontal inheritance common in
microbes and generalized to all forms of vertical, oblique and
horizontal transmission of genetic, epigenetic, parental, cultural
and ecological inheritances (Danchin et al. 2011, Danchin 2013).
The EES was developed to better understand the evolution of
complex phenotypic traits in nonhuman taxa, such as bird song,
tool use, and active allogenic ecosystem engineering that are not
readily explained without multigenerational feedbacks among
cultural, ecological, and/or genetic/epigenetic inheritances
(Danchin et al. 2011, Danchin 2013). Nevertheless, by building
ecological and cultural inheritance into evolutionary theory, the
TABLE 3. Recommended readings.
Core book (Odling-Smee et al. 2003c); review
Niche construction
for ecologists (Matthews et al. 2014)
theory
The Extended
Evolutionary
Synthesis
Cultural evolution
General review (Danchin et al. 2011), research
challenges (Danchin 2013)
Human
ultrasociality
Core book (Richerson and Boyd 2005); research
synthesis (Mesoudi et al. 2006); brief summary
(Castro and Toro 2010)
Evolution and human behavior (Laland and
Brown 2011)
Behavioral sciences (Tomasello et al. 2005);
anthropology (Hill et al. 2009)
Human niche
construction
Social learning and sociality (Sterelny 2011);
domestication (Smith 2012)
Social sciences
Societal types, Macrosociology (Nolan and
Lenski 2010)
Social Change, World Systems theory (ChaseDunn and Lerro 2013)
Human ecology theory (Butzer 1982);
environmental change (Redman 1999)
Global change (Steffen et al. 2007)
Terrestrial ecology (Ellis 2011)
Archaeology (Smith and Zeder 2013)
Paleontology (Barnosky 2014)
Geology (Zalasiewicz et al 2012)
Archaeology
Anthropocene
EES paves the way for major advances in understanding the
evolution and ecology of human societies. Though many species
exhibit cultural inheritance, which is defined as the transmission
of information across generations through social learning (i.e.
"learning from others"; Danchin et al. 2011), human capacity for
social learning and accumulating cultural inheritance across
generations is unsurpassed by any other species (Tomasello
1999). By bringing cultural, ecological and other inheritances
together within a single evolutionary model, the EES provides a
mechanistic framework for understanding the coupled evolution
of human cultural and ecological inheritance over generational
time.
One major prediction of the EES is that genetic and
epigenetic inheritances will tend to predominate in environments
that remain stable across generations and become less important
in transmitting phenotypic variation across generations as
environmental variation increases (Danchin 2013). As crossgenerational environmental variation increases, cultural and then
ecological inheritances become increasingly important modes of
phenotypic trait transmission, as traits that enhance phenotypic
plasticity are favored over the inheritance of traits adaptive only
under prior environmental conditions (Danchin 2013). Given
that ecosystem engineers generally reshape environmental
patterns and processes at levels that alter environmental
conditions experienced by later generations, the role of
ecological and cultural inheritance in the evolution of
phenotypic traits in ecosystem engineers would be expected to
increase, even more so in the case of culturally inherited traits
for ecosystem engineering, such as tool-use and domestication
(O’Brien and Laland 2012). To understand long-term
anthropogenic ecological changes caused by culturally
transmitted traits for ecosystem engineering, it is necessary to
understand how these cultural traits evolve, supported by
exceptional human capacities for social learning and sociality
that have enabled the emergence and evolution of sociocultural
systems of unprecedented scale, complexity and power.
Cultural evolution
Darwin used cultural analogies to explain biological
evolution (Darwin 1859, Mesoudi et al. 2004) and addressed
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human cultural evolution directly (Darwin 1871) helping to
spark the production of a superabundance of theory- much of it
incorrectly interpreting Darwin (Mesoudi et al. 2004, Laland and
Brown 2011). At least three major theoretical frameworks now
address human cultural evolution (Laland and Brown 2011):
sociobiology (Wilson 1975), gene-culture coevolution (CavalliSforza and Feldman 1981, Boyd and Richerson 1985, Durham
1991), gene-culture coevolution plus niche construction (closely
related to the EES; Laland et al. 2000), and a fourth framework,
less popular among evolutionary theorists, of “memetics”, with
“memes” serving as the cultural equivalent of genes (Dawkins
1976, Blackmore 1999, Laland and Brown 2011). Sociobiology
explains behavior genetically, avoiding cultural inheritance,
while noting that “the most spectacular cultural advances [of
humans] were impelled by the invention of new ways to control
the environment” (Wilson 1975). Memetics applies evolutionary
theory to socially learned information (“memes”), focuses
primarily on horizontal, not cross-generational transmission, and
generally ignores interactions with ecological processes
(Henrich et al. 2008). Gene-culture coevolution or “dualinheritance” theory explains the coupled evolution of genetic
and cultural traits, such as lactose tolerance in dairy farming
societies, without incorporating ecological inheritance (Laland
and Brown 2011).
Gene-culture coevolution plus niche construction integrates
genetic, cultural and ecological inheritances in a form capable of
explaining the evolution of cultural traits for ecosystem
engineering. The theory was developed by integrating geneculture coevolution (Cavalli-Sforza and Feldman 1981, Boyd
and Richerson 1985) with ecological inheritance (Laland et al.
2000, Mesoudi et al. 2006, Bonduriansky and Day 2009,
Mesoudi et al. 2013) to explain genetic changes in societies
engaged in ecosystem engineering, such as increased
frequencies of malaria resistance genes in rainforest yam
cultivating societies; an adaptive genetic response to malariacarrying mosquitos that are an indirect detrimental ecological
inheritance of yam cultivation in rainforests (Laland et al. 2000).
By considering this theory as a subset of the EES (setting aside
epigenetics and parental effects) in which genetic, cultural and
ecological inheritances interact in producing phenotypic
outcomes, we obtain a mechanistic basis for explaining crossgenerational changes in cultural niche construction: the
engineering of ecosystems by socially transmitted behaviors that
produce heritable adaptive benefits and/or detriments (Laland et
al. 2000, Laland et al. 2001, Mesoudi et al. 2006, Smith 2007a,
Bonduriansky and Day 2009, Boyd et al. 2011, Kendal et al.
2011, Laland and O’Brien 2012, O’Brien and Laland 2012,
Mesoudi et al. 2013).
Mechanisms of cultural evolution. - Culture is “the part of
phenotypic variation that is inherited socially (that is, learnt
from others)” (Danchin et al. 2011). Four criteria distinguish
whether a specific trait, or variant, is culturally inherited and
therefore capable of undergoing evolution by natural selection
(Danchin et al. 2011). First, a cultural trait must be socially
learned and not inherited by another transmission pathway or
acquired by individual (“asocial”) learning. Second, crossgenerational transmission of the trait must occur, generally from
older to younger generations (vertical or oblique transmission).
Third, organisms must exhibit the trait for sufficient time and in
such a way that others are capable of learning it. And fourth, the
trait must be expressed and selected for under differing
environmental conditions, because traits expressed or selected
for only within a single nonrepeating environmental condition
cannot be selected for across generations. By these criteria,
cultural inheritance is evident in many nonhuman animal species,
including the social learning of foraging strategies, tool use,
mate choice, and song dialects, though comprehensive evidence
meeting all four criteria is still lacking in nonhumans
(Tomasello et al. 1993, Danchin et al. 2011). In humans
however, evidence for cultural inheritance and cultural evolution
is overwhelming (Henrich and McElreath 2003, Laland and
Brown 2011, Mesoudi 2011, Whiten et al. 2011). Primary
among the cultural inheritances that humans have specialized in
is technology, defined here as cultural information enabling or
enhancing the behavioral capacity of organisms beyond their
biological capabilities, from the use of tools and the control of
fire, to the manufacture of advanced machines and computers
(Pfaffenberger 1992).
Macro- and micro-evolution of human cultural traits across
generations including toolmaking and other technologies has
been confirmed by mechanistic and empirical investigations
both across societies (macro-) and within societies and social
groups (micro-) (Basalla 1988, Henrich and McElreath 2003,
Mace and Holden 2005, Mesoudi et al. 2006, Bettinger 2009,
Boyd et al. 2011, Mace and Jordan 2011, Mesoudi 2011, Whiten
et al. 2011). That many human behavioral traits are cultural is
beyond question; individuals of different cultures readily acquire
and maintain cultural traits learned from each other (Laland and
Brown 2011). While the units of cultural inheritance remain
subject to ongoing scientific debates, it has been demonstrated
that the existence of cultural “replicators” analogous to genes
(e.g. memes) is not an absolute requirement for culture to be
inherited, only some form of copying behavior that leads to
some level of cross-generational transmission, whether vertical,
oblique or horizontal (Mesoudi et al. 2004, Henrich et al. 2008,
O'Brien et al. 2010).
For cultural traits to evolve by natural selection across human
generational time requires that these traits must vary within
generations, be transmissible across generations and cause
differential effects on fitness, such that the frequencies of
cultural traits vary over generations in response to selective
pressures. All of these properties are well confirmed for cultural
traits (Henrich and McElreath 2003, Mesoudi et al. 2006,
Mesoudi 2011, Whiten et al. 2011). The generation of novel and
variant cultural traits by asocial processes of innovation and trial
and error experimentation provides a source of cultural trait
variation analogous to the role of mutation in biological
evolution (Cavalli-Sforza and Feldman 1981, Laland et al. 2000,
Mesoudi and O'Brien 2008, Laland and Brown 2011). Cultural
traits are transmitted with high fidelity both vertically and
obliquely by human behaviors that include the copying of
common cultural traits (conforming), the imitation of highperforming cultural traits, and the teaching of traits to progeny
and nonkin individuals (Cavalli-Sforza and Feldman 1981, Boyd
and Richerson 1985, Tomasello et al. 1993, Richerson and Boyd
1998, Eerkens and Lipo 2007, Laland et al. 2007, Laland et al.
2010, Chudek and Henrich 2011, Laland and Brown 2011,
Lewis and Laland 2012). Long-term trends in cultural traits also
show patterns analogous to those observed for genetic traits
(Cavalli-Sforza and Feldman 1981, Mesoudi et al. 2006, Laland
and Brown 2011; Table 6.1 ), including long-term phylogenetic
trends in languages and material culture, such as arrowheads
(Table 1; (Mace and Holden 2005, Eerkens and Lipo 2007,
O'Brien et al. 2010, Gray et al. 2011, Mace and Jordan 2011,
Shennan 2011a), the accumulation of cultural information,
diversity and complexity (Mesoudi et al. 2004, Foley and
Mirazón Lahr 2011), convergent patterns such as writing and
domestication (Mesoudi et al. 2004), extinctions (Mesoudi et al.
2006), and the persistence of vestigial patterns like the
QWERTY keyboard (Basalla 1988).
Studies of African tribes have demonstrated that most human
cultural trait transmission may be vertical, not horizontal
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(Guglielmino et al. 1995). Nevertheless, high levels of
horizontal cultural trait transmission have been observed,
especially in larger scale societies, and frequencies of cultural
traits can vary dramatically within generations and within and
across human social groups and societies (Mesoudi et al. 2006,
Henrich et al. 2008, Laland and Brown 2011, Godfrey-Smith
2012). Selection processes acting on horizontally transmitted
cultural traits within the span of a human generation can enable
cultural traits to evolve far more rapidly than genetic traits
(Henrich and McElreath 2003). While the horizontal
transmission of genetic traits is also important in
microorganisms and in viruses, rates of horizontal transmission
of human cultural traits and their selection at group and
population levels adds new levels of complexity to evolutionary
patterns and processes (Henrich et al. 2008, Laland and Brown
2011, Godfrey-Smith 2012). For example, horizontal exchanges
of words and technologies across societies ("cultural borrowing")
challenges phylogenetic analysis of human cultural history,
blending together the branches of otherwise tree-like cultural
lineages (Gray et al. 2010). And that is only the beginning of the
evolutionary complexities introduced by cultural inheritance.
Cultural inheritances may amalgamate other cultural
inheritances, for example the knowledge needed to manufacture
a tool, which archaeologists conceive of as “recipes” combining
the preparation of raw materials, tool construction, use, repair
and maintenance (Mesoudi and O’brien 2008). Even greater
complexity is added by the potential for cultural inheritance to
alter the adaptive fitness of entire groups or societies, with the
result that natural selection has the potential to act at the level of
entire competing groups of kin and/or nonkin individuals
through the adaptive advantages of social foraging, collective
strategies for warfare and defense, material exchange, and even
the collaborative advantages of language (Boyd and Richerson
1985, Henrich 2004, Bowles 2006, 2009, Laland et al. 2010,
Apicella et al. 2012, Rand and Nowak 2014). Given the clear
adaptive fitness benefits for individuals, groups, and individuals
within groups engaging in collaborative social behaviors, it is
not surprising that selective processes acting at all of these levels
are implicated in the evolution of the unprecedented social
behaviors present in humans (Boyd and Richerson 1985,
Chudek and Henrich 2011, Rand and Nowak 2014).
Human ultrasociality and the human sociocultural niche
Two exceptional patterns of human social behavior are
needed to explain the emergence of humans as a global force
transforming the biosphere. The first is the unrivalled capacity
of humans to transmit information by social learning,
exemplified by human use of language (Tomasello 1999, Hill et
al. 2009, Pinker 2010, Dean et al. 2012, Morgan et al. 2015).
Compared even with close relatives, including Chimpanzees and
Apes, humans are far more capable of social learning across
both related and nonrelated individuals and especially across
generations (Tomasello 1999, Pinker 2010, Laland and Brown
2011, Dean et al. 2012, Morgan et al. 2015). Second, humans
have unparalleled capacity to form, sustain, and depend for
survival on complex nonkin relationships, easily marking
humans as the most ultrasocial species on Earth (Campbell 1983,
Richerson and Boyd 1998, Fehr and Fischbacher 2003,
Tomasello et al. 2005, Boyd and Richerson 2009, Hill et al.
2009, Boyd et al. 2011, Apicella et al. 2012, Turchin et al. 2013,
Tomasello 2014).
There is increasing evidence that some aspects of the
exceptional human capacity for social learning, including use of
languages, is supported by genetic traits, some of which have
been identified recently as specific to Homo sapiens, including a
variant of FoxP2 (a protein involved in brain development and
language) among others (Enard et al. 2002, Tomasello et al.
2005, Laland et al. 2010, Dean et al. 2012, Fisher and Ridley
2013, Pääbo 2014). Gene-culture coevolution is strongly
implicated in the early evolution of human genetic traits
supporting both social learning and ultrasociality (Boyd and
Richerson 1985, Laland et al. 2010, Chudek and Henrich 2011,
Gintis 2011, House et al. 2013), and molecular genetic
comparisons among extant humans, extinct hominins, and other
close relatives is enabling unprecedented breakthroughs in
understanding the evolution of human genetic traits relating to
social learning and ultrasociality over thousands to millions of
years (Pääbo 2014).
Yet the role of genetics in explaining the emergence of
modern human behaviors approximately 60,000 years ago in
Africa has remained controversial (Henshilwood and Marean
2003, Mellars 2005, Nowell 2010, Fisher and Ridley 2013,
Klein 2013, Sterelny 2014). Some experts argue that a genetic
mutation or other major genetic change is required to explain the
relatively sudden and widespread emergence of the suite of
modern human behaviors (Klein 2013), which include symbolic
inscriptions, lithic projectile point weapons, personal ornaments,
rapid changes in tools and technologies, more highly structured
settlements and long-distance trade (Mellars 2005, Hill et al.
2009, Nowell 2010). Others explain this based on the need to
adapt to rapid environmental changes or heterogeneous
environmental conditions (Mellars 2005, Stiner and Kuhn 2006,
Potts 2012), to changes in social conditions conducive to social
learning (Sterelny 2011), including demographic shifts toward
higher population densities enabling greater accumulation of
cultural information (Shennan 2001, Powell et al. 2009, Derex et
al. 2013), the emergence of exchange-based economies
(Sterelny 2014), increased intergroup conflict or cooperation
(Mellars 2005, Stiner and Kuhn 2006), and by other factors and
combinations of factors (Powell et al. 2009).
Despite disagreements on the role of genetics in the
emergence of behavioral modernity, there is broad consensus
that the behaviorally modern human populations that spread out
of Africa more than 50,000 years ago possessed genetic
capacities for social learning and sociality that were functionally
equivalent to those of human populations today (Sterelny 2011,
Pääbo 2014). Human genetic evolution continues to respond to
ecological pressures including diseases and environmental
conditions, some of which have resulted from human sociality,
such as high population densities, trading networks, and
engineered environments (Laland et al. 2010, Richerson et al.
2010, Rendell et al. 2011). Nevertheless, genetic changes cannot
explain long-term changes in modern human behaviors that
support social learning and sociality or their effects on the
structural organization of human societies.
Human cultural traits for sociality have evolved over the long
term. For example, common social behaviors within huntergatherer societies, such as high degrees of resource sharing, are
incompatible with those common in industrial societies
(Richerson and Boyd 1998, Richerson and Boyd 1999, Henrich
et al. 2001, Smith et al. 2010a). Human social institutions,
including formal and informal rules, conventions, and other
forms of socially learned information that regulate and structure
human relationships within societies and sometimes across
societies have also changed significantly over the long-term and
also show great diversity across and sometimes within societies
(Richerson and Boyd 1999, Hodgson 2002, Boyd and Richerson
2008, Kaplan et al. 2009, Pinker 2010, Henrich 2015).
Cultural traits are what enable behaviorally modern humans
to sustain themselves and their progeny within social groups and
societies and cultural traits also produce and sustain the social
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organization of these groups and societies; both are the product
of ongoing cultural evolution (Richerson and Boyd 1999,
Henrich et al. 2001, Hodgson 2002, Hill et al. 2009, Kaplan et al.
2009, Smith et al. 2010a, Boyd et al. 2011, Sterelny 2011). As a
result, the human niche is largely sociocultural; defined within
the patterns and processes of sociocultural systems together with
the altered ecosystems that sustain them. Further, the human
niche is also dynamic in response to changes in social
organization brought about both by cultural evolution and the
environmental changes caused by cultural niche construction.
Human agency and intentionality
Sociocultural systems emerge and reproduce through the
interactions of individual humans and social groups as agents
deciding and acting within culturally inherited social structures
("structuration theory"; Giddens 2013). While the actions of
individuals within sociocultural systems can depend largely on
culturally inherited traits (Richerson and Boyd 1999, Henrich et
al. 2001), individuals and groups choose among and adopt
cultural traits in different ways, such as copying the most
common traits in a population or group versus copying those of
high status individuals or groups or even low status groups,
yielding substantial variance and unpredictability in individual,
group and societal behavior (Henrich 2001, Macy and Willer
2002, Brown et al. 2011a, Chudek and Henrich 2011, Gelfand et
al. 2011, Wolf and Krause 2014). Though human capacity for
cooperative behavior is exceptional, so is human cognitive
capacity and behavioral flexibility (Roth and Dicke 2005,
Brown et al. 2011a). As a result, human agency and
individuality of choice are key determinants of the emergent
behavior of sociocultural systems and groups, such that
culturally inherited social structures and behaviors alone are
incapable of fully predicting individual, group, or societal
behavior (Macy and Willer 2002, Brown et al. 2011a, Gelfand et
al. 2011, Smith 2013b). Further, humans have a capacity for
shared intentionality not present in any other species (fourth
order intentionality); the ability to interpret the intentions of
others and to act cooperatively with these intentions, enabling
individual choices to scale up to larger group decisions (Dunbar
1998, Tomasello et al. 2005, Dean et al. 2012). This unrivalled
ability to act intentionally and cooperatively adds another
dimension to the consequences of human agency: intended vs.
unintended consequences, together with social responsibility for
these consequences. For example, ecosystem engineering by
tilling soils might yield a combination of its intended beneficial
direct consequence, enhanced food supply, together with
unintended detrimental direct consequences, such as a long-term
decline in soil fertility, and unintended indirect detrimental
consequences, such as water pollution with sediments affecting
human populations downstream.
The ratchet effect and runaway cultural niche construction
Though human individuals and social groups can act
intentionally to alter sociocultural systems and ecosystems,
long-term, cross-generational changes in sociocultural systems
and human engineered ecosystems are shaped largely by
processes of cultural evolution. The overwhelming importance
of cultural over genetic traits in human niche construction is
explained partly by the observation that cultural traits can evolve
much faster than genetic traits. While cultural traits have been
observed to evolve slowly, at rates similar to those of genetic
traits, for example the evolution of technologies for stone tool
manufacture by early hominins (Laland et al. 2000, Nolan and
Lenski 2010, Table 5.1), the hypothesis that cultural traits can
evolve far more rapidly than genetic traits has had wide support
since it was first proposed by Darwin (Mesoudi et al. 2004,
Richerson and Boyd 2005, Eerkens and Lipo 2007). Horizontal
transmission and selection among cultural traits within a single
human generation is one process that can enable cultural traits to
evolve faster than genetic traits (Henrich and McElreath 2003).
The “ratchet effect” is a further cause of rapid cultural evolution,
as cultural traits can build upon and combine earlier cultural
traits to produce increasingly complex and powerful cultural
traits such as cultural institutions (e.g. languages, legal systems)
and advanced technologies (projectile weapons, the automobile)
that enable cultural accumulation to accelerate across
generations (Tomasello 1999, Laland et al. 2000, Dean et al.
2014).
Perhaps the most powerful mechanism supporting rapid
cultural change is “runaway” cultural evolution, in which
changes in culture must be adapted to by further changes in
culture, which in turn require additional cultural adaptations,
generating accelerating rates of cultural change across
generations (Boyd and Richerson 1985, Laland et al. 2000,
Richerson and Boyd 2005). This runaway effect can even
increase the frequency of maladaptive cultural traits, such as
when individuals copy the prestige-seeking behaviors of
influential or successful members of their society, such as costly
adornments, grave monuments, and other forms of conspicuous
consumption (Boyd and Richerson 1985, Boyd et al. 2011).
Rates of evolution of cultural traits for ecosystem engineering
(cultural niche construction) appear to have become so rapid that
they have overwhelmed natural selection for human genetic
adaptations to environments (Laland et al. 2000, Laland et al.
2001). The central mechanism proposed to explain this is
runaway cultural niche construction, in which socially learned
traits for ecosystem engineering cause environmental changes
that must be adapted to by additional cultural traits (Rendell et al.
2011, Laland and O’Brien 2012). For example, tillage of soils to
produce crops reduces soil productivity over time, requiring
fallowing, manuring or other cultural practices to maintain the
productivity of soils across generations. The use of antibiotics
and the development of antibiotic resistance, requiring further
antibiotic development is another. By processes of runaway
cultural niche construction combined with the ratchet effect,
cultural traits for niche construction tend to become increasingly
adaptive, complex and powerful across generations, the
evolution of these traits is accelerated, and populations become
more and more dependent on cultural traits for ecosystem
engineering to sustain themselves (Laland et al. 2007, Rendell et
al. 2011).
There are further explanations for increasing human
dependence on cultural niche construction. Socially learned
traits for ecosystem engineering have the potential to support
larger human populations, and larger populations have the
potential for more rapid cultural evolution, especially in small
scale societies (Shennan 2001, Kline and Boyd 2010, Derex et al.
2013). Population pressures in themselves can drive demand for
cultural adaptations to larger and denser populations, a potential
explanation for observed relationships between cultural
complexity and population among hunter gatherers (Keeley
1988, Powell et al. 2009, Muthukrishna et al. 2014, Vegvari and
Foley 2014). But no matter what the mechanisms, there is no
question that socially learned practices of ecosystem engineering
have changed dramatically over the past 50,000 years and
increased in their transformative capacity and complexity
together with the diversity, scale, and complexity of human
sociocultural systems and their transformations of the biosphere.
Human sociocultural niche construction
By
Page 10 of 40
combining
cultural
niche
construction,
culturally
Ellis, E.C. 2015. Ecology in an Anthropogenic Biosphere. Ecological Monographs in press doi.org: 10.1890/14-2274.1 [FORMATTED PREPRINT]
mediated social organization, and cultural evolution into a single
theory of sociocultural niche construction (SNC; Tables 1, 2, 3,
4), the observation of dramatic long-term changes in and
diversification of the human niche can be explained, together
with the capacity of human societies to transform the biosphere.
Just as species engage in cultural niche construction when their
socially-learned behaviors alter environments in ways that
produce heritable adaptive consequences, humans engage in
sociocultural niche construction when their socially learned
behaviors are enacted socially, altering both the organization of
their societies and the environments that sustain them in ways
that produce heritable adaptive consequences. As with cultural
niche construction, sociocultural niche construction, unfolding
across generations, tends to cause increasing dependence on
cultural traits for survival and reproduction, including
technology, socially learned subsistence strategies for
cooperative ecosystem engineering and nonkin subsistence
exchange; the socially mediated exchange of materials, labor,
and information to meet subsistence needs (Table 1). To the
extent that subsistence strategies for cooperative ecosystem
engineering and subsistence exchange are socially learned and
socially enacted (dependent on cooperative efforts across groups
or societies rather than individuals acting alone), these are
“subsistence regimes” (Table 1), or cooperative processes that
sustain social groups and societies that cannot be implemented
by individuals alone.
Evidence for human sociocultural niche construction is
presented in Table 4. As processes of sociocultural niche
construction have evolved across generations, the human niche
has broadened dramatically, diversified within and across social
groups and societies, and has become increasingly social, such
that most human individuals have come to depend more and
more on complex networks of social interaction and nonkin
subsistence exchange for their survival and do this at increasing
spatial scales (Hill et al. 2009, Kaplan et al. 2009). Increasing
dependence on nonkin subsistence exchange networks has in
turn enabled human societies to become increasingly specialized,
complex, and hierarchical (Hill et al. 2009, Nolan and Lenski
2010, Chase-Dunn and Lerro 2013), with individuals specialized
in different socially learned productive capacities cooperating
with unrelated and often unknown individuals through longdistance exchange networks to accomplish complex tasks. One
example is the production of shell-bead garments, which require
shell harvest and preparation in coastal areas, long-distance
TABLE 4. Evidence for human sociocultural niche construction (SNC).
1. The human ecological niche has changed dramatically over time. Rates of change in the human niche appear to be more rapid than possible by
biological evolution, and these rates of change appear to be generally increasing over time.
2. The human ecological niche is not predictable from human biology.
2.1 Human genetic variation does not account for variation in human subsistence strategies or habitat preferences, over space or time.
2.2 Genetically similar, demographically equivalent human individuals and populations within the same environment can engage in extremely different
livelihood strategies based on socially learned behaviors.
2.3 Human use of biotic and abiotic resources and alteration of ecosystems is not strictly density dependent. Profoundly different demands and effects are
observed at similar population densities in similar environments. These differences are generally associated with differences in sociocultural systems.
3. The human ecological niche is broader and more diverse than that of any other species.
3.1 Humans live under a broader range of environmental conditions than any other multicellular species and are more widely distributed across the Earth.
3.2 Humans utilize a wider range of biotic, abiotic, and energy resources than any other species.
3.3 Humans engineer ecosystems and apply selective pressures to species by a larger number of diverse practices and at levels and scales greater than that
of any other species.
4. Different human societies and social groups transform ecology in a variety of different ways.
4.1 When one socioculturally distinct group displaces another across a given region, ecological pattern and process generally tends to change as well.
4.2 Human populations and their effects can be dynamic in stable environments, stable in dynamic environments and vice versa.
4.2 Human populations and their effects can be heterogeneous across homogeneous environments or homogeneous across heterogeneous environments
and may or may not follow pre-existing environmental patterns.
5. Humans depend on nonkin subsistence exchange far more than any other species.
5.1 Humans engage in prosocial nonkin interactions at the expense of individual fitness. This trait shows both genetic and cultural inheritance.
5.2 Humans regularly exchange materials, biota, energy and information across extensive, complex and dynamic nonkin networks. These social exchanges
may now be daily, global and even extraterrestrial.
5.3 Human populations can depend entirely on nonkin exchanges of food and other necessary material and energy resources with other populations for
survival.
5.4 Nonkin exchange can sustain the long-term growth and development of human populations, decoupling them from the use of local ecological, material
and energy resources.
5.5 Nonkin exchange can facilitate the utilization and accumulation of resources at much higher levels than possible by utilizing local resources.
5.6 Subsistence and other demands of human populations can exert ecological effects over long distances by means of nonkin exchange relations
(trade/telecoupling).
5,7 The relative scale and distance of trade/telecoupling tends to increase over time.
6. Socially learned subsistence strategies for ecosystem engineering and nonkin exchange exhibit cross-generational heritability and competition.
6.1 Selection for desired traits has been applied to some species for millennia, yielding domesticates with little resemblance to ancestral species that are
poorly adapted to non-human habitats (e.g. maize, wheat, dogs).
6.2 Anthropogenic ecosystems requiring high levels of human maintenance have been sustained for centuries to millennia (e.g. rice paddy irrigation
systems, cities).
6.3 Socially learned subsistence strategies for ecosystem engineering and specialized exchange including agriculture and craft industries have been
transmitted with high fidelity across large numbers of generations and disparate populations and societies.
6.4 Variants of socially learned subsistence strategies, including technologies, tools, and institutions governing social exchange show long-term changes in
frequencies and phylogenetic relationships demonstrating that competition, selection and accumulation of variations in these strategies have occurred
over generations.
7. The scale, structural complexity, specialization, and ecological transformative capacity of human societies and their subsistence strategies vary
tremendously and have tended to increase over the long-term.
8. Human alteration of local and global environments is now greater than that of any other multicellular species.
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trade (social exchange), the production of hides or textiles in an
upland area, and their integration by a skilled bead worker in
another area. Specialization and exchange in subsistence
regimes have made it possible for human individuals to subsist
apart from any direct interactions with ecosystems (though not
without indirect interactions, or telecoupling; Table 1), with all
subsistence needs met through exchange networks of
subsistence producers (i.e. farmers, fisherman), processors (food
preparation), providers (traders) and potentially many more
specialists (tool makers, irrigation experts, bankers) in complex
and dynamic subsistence supply chains (“subsistence webs”)
inviting further study as “socio-trophic relations”.
As specialization and exchange have increased, human
interactions with ecosystems have increasingly become societal
interactions based on subsistence regimes enacted by ever larger
groups of cooperating specialized individuals and groups guided
by accumulated cultural inheritances of social organization,
social exchange and technological capacities far beyond those of
any human individual. The human sociocultural niche is a
function of the accumulated cultural inheritance of societies and
is therefore the product of and subject to cultural evolution (Hill
et al. 2009, Kaplan et al. 2009, Boyd et al. 2011).
SOCIAL CHANGE: SOCIETAL SCALE, COMPLEXITY,
TECHNOLOGY AND INTERACTION
As with biological evolution, there is no simple progressive
pathway describing the rise, diversification and extinction of
societies, or the observed tendency towards increasing scale and
complexity. Some of the earliest forms of sociocultural systems,
such as hunter gatherer societies, are remarkably complex and
have endured to the present day, sometimes in the face of
pressures from larger scale societies (Marlowe 2005). Yet, as
with the convergent evolution of similar phenotypes across taxa,
human sociocultural systems, while tremendously diverse,
complex and heterogeneous, also show some converging
patterns over time. These basic patterns have been categorized
by social scientists into societal types based on major
differences in their primary subsistence regimes, as described in
Table 5 (Lenski 1966, Nolan and Lenski 2010).
To understand the emergence and divergence of sociocultural
systems over time, it must first be noted that the data available
in Table 5 are neither complete nor exclusive- some societal
types are missing, societies regularly intermingle to generate
hybrid forms, and the patterns of a given societal type are
generally not shared by all societies of the same type. Still, some
general patterns of long-term sociocultural change are evident
when societal types are ordered in relation to the order of their
appearance (Table 5; Lenski 1966, Nolan and Lenski 2010).
Over time, the scale of societies has increased by more than 5
orders of magnitude from the small-scale societies of hunter
gatherers to large-scale industrial societies, and as societal scales
have increased, so have societal complexity, specialization, and
technological capacity, along with inequality among individuals
within societies in the distribution of resources, social power
(the capacity to act independently), and access to environments
(Mulder et al. 2009, Nolan and Lenski 2010, Smith et al. 2010b,
Chase-Dunn and Lerro 2013).
Lenski (1966) introduced a theory of sociocultural evolution
to explain these patterns in which the social organization of
societies, including their institutions, beliefs, complexity and
degree of specialization are seen as responses to the population
sizes and densities sustainable by their primary subsistence
technologies “which define the limits of what is possible for a
society” together with the accumulation of these and other
cultural inheritances over time through cultural evolution (Nolan
and Lenski 2010). In other words, technological innovation and
its accumulation through cultural inheritance are theorized as the
ultimate drivers of long-term social change. Lenski’s theory also
holds that societies evolve over the long-term both through
internal processes and by competition, exchange and other
interactions among societies within a “world system” (Table 1).
While Lenski’s (1966) theory of sociocultural evolution has not
garnered mainstream support, his identification of long-term
societal trends and theory of world systems remain core
elements of contemporary sociology and anthropology (ChaseDunn 2006, Nolan and Lenski 2010, Hall et al. 2011, ChaseDunn and Lerro 2013).
Of the many theories explaining long-term societal change,
all tend to incorporate relationships among human demographics,
subsistence technologies and other cultural inheritances (cultural
complexity), social organization and institutions (social
complexity and inequality), and some also include energy use
(White 1959, Flannery 1972, Butzer 1982, Redman 1999,
Redman et al. 2004, Sanderson 2006, Tainter 2006b, Abrutyn
and Lawrence 2010, Nolan and Lenski 2010, Tainter 2011,
Butzer 2012, Chase-Dunn and Lerro 2013, Fischer-Kowalski et
al. 2014). Another agreement across theories is that most change
tends to be gradual, punctuated by relatively abrupt societal
transitions or regime shifts (Geels 2002), including episodes of
relatively rapid growth and societal collapse, in which
populations, technological capacity (linked to energy use),
cultural complexity, and social organization tend to change
together (Tainter 2006b, 2011). The degree to which innovations
in technology in themselves determine the long-term patterns of
sociocultural change has long been debated without resolution,
with theories ranging from “technological determinism” (Smith
and Marx 1994) to “social construction of technology” (Bijker et
al. 1987) to coevolutionary models of social and technological
change (Basalla 1988, Nelson 1994, Redman 1999, Geels 2002,
Geels and Schot 2007, Hodder 2011, Brynjolfsson and McAfee
2014, Haff 2014, Morgan et al. 2015). For a variety of reasons,
especially the challenge of establishing appropriate models,
causal relations among population, technology, cultural
inheritances and social complexity have yet to be established
scientifically (Hedström and Ylikoski 2010). Nevertheless, their
tight linkage across societal scales and their tendency to change
together in regime shifts does imply that these are
mechanistically coupled in multicausal relationships.
Interactions among societies also help to explain general
long-term trends of societal change. Analogous to competition
among individuals, competition among societies, including
warfare, should select for cultural traits that enhance adaptive
fitness in the face of intersocietal competition, such as weaponry,
strategies for warfare, raiding, defense, and avoidance, together
with larger scales of social organization (Nolan and Lenski 2010,
Chase-Dunn and Lerro 2013, Turchin et al. 2013, Scott 2014).
That larger societal scales can become an adaptive advantage
over smaller scales is summarized by the “law of cultural
dominance”, which holds that “larger scale societies tend to
destroy or radically alter the cultures of smaller scale societies”
(Sahlins et al. 1960, Nolan and Lenski 2010, Chase-Dunn and
Lerro 2013). Beyond its strong empirical support and the
obvious advantage of numbers, the law of cultural dominance
has also been explained by the theory that larger populations
have greater capacity for cultural innovation, cultural
accumulation, and stronger social organization (Powell et al.
2009, Nolan and Lenski 2010, Chase-Dunn and Lerro 2013,
Turchin et al. 2013).
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TABLE 5. Human sociocultural systems classified by primary subsistence regime, in order of historical emergence, based on Lenski (1966), updated by
Nolan and Lenski (2010) using data from Murdock and White (1969, 2006). Hybrid systems are common (e.g. “industrializing agrarian” = industrial +
agrarian), especially in larger and more complex societies, Fishing, maritime, and other less common societies not included. “nd” = no data
Sociocultural
system
Subsistence
regime1
Technological
innovations1
Cultural &
institutional
innovations2
Hunter
Gatherer
hunting,
foraging
land clearing using
fire, social hunting,
food processing &
cooking, projectiles,
ceramics
Simple
Horticultural
long fallow
shifting
cultivation
Herding
Social
complexity4
3
Scale
Specialization
5
Energy6
(total|food)
-1
Density7
(median
population)
(% complex)
(% specialized)
(GJ person
yr -1)
languages,
barter,
permanent
settlements,
tribes
40
0
0
8|5
2
domestication,
tillage (hoe)
land ownership,
trade, villages
1500
1
2
nd | 5
35
nomadic
pastoralism
horse warfare
extended trading
networks
6000
nd
9
nd | 6
nd
Advanced
Horticultural
short fallow
shifting
cultivation,
conquest
nonferrous metals
(weapons),
manuring, terracing
warfare/raiding,
chiefdoms
5000
7
22
nd | 5
110
Simple
Agrarian
continuous
subsistence
agriculture,
handicrafts
plow, animal
traction
taxation, writing,
numeracy, cities,
states
18 | 6
>250
Advanced
Agrarian
subsistence &
commercial
agriculture,
specialized
crafts
iron (tools),
irrigation, roads,
printing, regional
trade
coined money,
empires
40 | 9
urban
commercial
agriculture,
manufacturing
fossil energy,
synthetics, rapid
bulk transport,
telecommunication
capitalist states,
banking, global
trade, science
>107
100
100
118 |11
“
commercial
agriculture,
services
nonbiological
energy, internet,
genetic engineering,
robotics
international
governance,
global peer
exchange
>109
100
100
350 | 15
“
Industrial
Post-Industrial
>105
51
(persons
km-2)
34
1
Based on Nolan and Lenski (2010) Table 4.1.
Cultural system where earliest form of cultural and institutional innovations have been observed (Chase-Dunn and Lerro 2013).
Median population size of societies, by type of society, from Nolan and Lenski (2010) Table 4.2..
4
Percentage of societies having complex status systems, from Nolan and Lenski (2010) Figure 4.3.
5
Average frequency of craft specialization across societies, by type of society, from Nolan and Lenski (2010) Table 4.3.
6
Approximate total and food energy consumption per capita from Table 6.1 in Christian and McNeill (2004). Total energy =home + agriculture +
commerce + industry + transport. Food energy = food + animal feed. Industrial estimates ca.1850.
7
Median population density, by type of society, from Table 6.1 in Nolan and Lenski (2010). “Urban” refers to societies in which high density urban
populations are spatially distant from the lower density agricultural lands that sustain them (decoupling).
2
3
Two other elements of intersocietal interaction are important,
subsistence exchange and cultural exchange (Chase-Dunn and
Lerro 2013). Starting with the first societies of behaviorally
modern humans, exchange through long-distance trade of
prestige goods is evident, such as shell beads (Mellars 2006),
and this exchange has grown dramatically in bulk and extent as
exchange of subsistence goods developed, with an early
example being the exchange of wheat from southern European
farmers to northern European hunter gatherers 8000 years ago
(Chase-Dunn and Lerro 2013, Smith et al. 2015). The horizontal
exchange of cultural information and populations among
societies has likely also been a feature of behaviorally modern
human societies since the beginning (Bellwood 2001, Mesoudi
et al. 2006, Skoglund et al. 2012, Chase-Dunn and Lerro 2013,
Barceló et al. 2014). These processes of material and cultural
exchange and the intermingling of populations and societies
through warfare and emigration have, together with the internal
processes of societies, sustained a long-term trend of increasing
societal scales, complexities, diversification, interconnection,
the accumulation of technology and other cultural inheritances,
and ultimately, the emergence of a diverse and complex globalscale world system (Chase-Dunn and Lerro 2013).
AN EVOLUTIONARY THEORY OF
LONG-TERM ANTHROECOLOGICAL CHANGE
By weaving together existing theories of ecosystem
engineering, niche construction, inclusive inheritance, cultural
evolution, ultrasociality, and social change we have the basis for
a theory explaining the long-term upscaling of human societies
and their unprecedented capacity to transform the biosphere
through long-term changes in human sociocultural niche
construction (Tables 1-3). The next step is to integrate these
within an evolutionary framework in which sociocultural
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systems, ecosystems, and sociocultural niche construction are
coupled with their cultural, ecological and material inheritances,
as “anthroecosystems” (FIG 2A). Material inheritances represent
the heritable adaptive benefits of artificial materials and
constructs, such as buildings, roads, and pollutants that are
produced exclusively by behaviorally modern human societies
and are incapable of forming by natural processes.
In the anthroecosystem framework, sociocultural systems and
the biota within ecosystems coevolve through the sustained
direct interactions of sociocultural systems and ecosystems over
human generational time (FIG 2A). Anthroecosystems are
formed of these systems and their cross-generational interactions,
and change through processes of natural selection acting on their
cultural, material, and ecological inheritances, with inheritances
conferring adaptive benefits to individuals, groups and societies
being selected for, and those producing detriments, against. In
such a way, anthroecosystems change through evolutionary
processes acting on sociocultural niche construction over the
course of human generations, accumulating, losing, and
combining cultural, material, and ecological inheritances
through gradual processes of selection, accumulation, attrition
and recombination, and also more rapidly by regime shifts in
subsistence regimes and social organization (FIG 2B).
Patterns of long-term change in
sociocultural niche construction
By applying the anthroecosystem framework (FIG 2) we may
examine major regime shifts in sociocultural niche construction
associated with major societal transitions (Table 5) in terms of
their relative cultural, material, ecological, and human genetic
inheritances as depicted in FIG 3A and can also be related to
long-term changes in ecosystem transformation and nonhuman
energy use (FIG 3B). Examples of subsistence regimes
producing the cultural, material and ecological inheritances in
FIG 3A are described in more detail with references in Table 6.
The emergence of anatomically modern Homo sapiens about
200 ka BP is depicted at far left in FIG 3, highlighting that
control of fire, meat-eating, rudimentary cooking, and the
manufacture of stone tools were already established cultural
traits of multiple hominin species long before the emergence of
our species (Ambrose 2001, Antón et al. 2014). A global
timeline of changes in human societies, populations and
ecosystem transformation is presented in FIG 4.
Hunter gatherers. - The rise of behaviorally modern human
populations in Africa >50 ka BP is associated with a major
expansion in the adaptive role of cultural inheritance as a wide
array of novel subsistence regimes emerged then, including
landscape modification for hunting, use of lithic projectile point
weapons for hunting, symbolic and aesthetic expression
(symbolic markings, cave paintings, ochre, beads), increasingly
complex and standardized tools made from a widened variety of
materials, and likely the use of languages to enhance
cooperation in foraging, trade and other social exchange and
social organization (Sterelny 2014). Genetic inheritance in
hominins narrows through the extinction of Homo
neanderthalensis, most likely as a result of competition and
FIG. 2. Conceptual model of (A) an anthroecosystem combining
sociocultural and ecological systems through heritable and pathdependent interactions, and (B) long-term, cross-generational changes
in anthroecosystems caused by sociocultural niche construction both
through gradual variations in inheritances and by regime shifts caused
by novel and transformative inheritances and combinations of
inheritances. Regime shift illustrated here depicts new trading system
(cultural + material inheritance) + facilitated species invasion (orange
dot) + new biotic interactions (arrows). Widening purple bar depicts
the increasing role of sociocultural niche construction in shaping
anthroecosystem structure and function. Path dependent abiotic change
is depicted her by erosive reshaping of brown landform.
displacement by larger populations of behaviorally modern
humans (Klein 2009). More importantly, runaway sociocultural
niche construction begins, with cultural and ecological
inheritances surpassing genetic traits in adapting to the
environmental alterations and social challenges of living in
culturally complex hunter gatherer societies (Sterelny 2014).
The adaptive benefits of material inheritances also begin to
accumulate through the manufacture and intergenerational
exchange of increasingly labor-intensive and complex
ornaments, tools, clothing and other material goods.
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FIG. 3. Conceptual model of regime shifts in human sociocultural niche construction across major types of sociocultural systems (Table 5). All Y axes
indicate relative, not absolute, changes. (A) Increasing scales of sociocultural niche construction (widening purple bar); also a proxy for societal scale,
median population size and degree of specialization of individuals within societies (Table 5) and their relative production of cultural, ecological,
material and genetic inheritance (relative heights of pink, gray and green bars). Nonhuman genetic inheritances are incorporated within ecological
inheritance. (B) Relative per capita energy expenditure and per capita ecosystem transformation in terms of relative anthrome area used across different
types of sociocultural systems (anthrome levels; legend same as in FIG 1B).
Before the Pleistocene had ended and the Holocene began,
behaviorally modern human hunter gatherer societies became
established on every continent, initiating the global role of
sociocultural niche construction as a force transforming the
biosphere through early forms of cooperative ecosystem
engineering and other socially learned and implemented
subsistence regimes (Kirch 2005, Doughty 2013, Ellis et al.
2013b). Early hunter gatherers introduced anthropogenic fire
regimes across the continents, both unintentionally, and with the
intention to create and maintain open landscapes and early
successional ecosystems to enhance their success in hunting and
foraging (Cronon 1983, Grayson 2001, Bird et al. 2005,
Bowman et al. 2011, Rowley-Conwy and Layton 2011, Smith
2011, Ellis et al. 2013b). More complex and sedentary hunter
gatherer societies developed larger populations, a degree of
social inequality and hierarchy (Ames 2007, Shennan 2011b),
and some developed ceramic technologies, the first artificial
minerals produced by humans (Craig et al. 2013). Larger
populations put greater pressure on plant and animal resources,
helping to drive megafauna extinct (Barnosky 2008) and
requiring the broadening of hunting and foraging strategies to
obtain adequate nutrition once preferred prey and plant
resources became rare (Stiner 2001, Zeder 2012). Technologies
for food processing, including grinding and boiling, increased
the nutrients extractable from plant and animal foods, boosting
food returns from limited land and potentially making small
seeds and tubers worth exploiting at high levels for the first time,
putting them on course to later domestication (Fuller et al. 2011,
Wollstonecroft 2011).
Sedentary populations of complex hunter gatherers,
especially those occupying the most productive parts of
landscapes, began to propagate desirable species of plants and
animals by a wide array of pre- and proto-agricultural niche
construction strategies (Price and Bar-Yosef 2011, Smith 2011,
2012). By the early Holocene, most human populations likely
lived in societies that had adapted to denser populations by
processes of sociocultural niche construction that boosted the
productivity of land through ecosystem engineering (Smith
2007b, 2011) and an array of other subsistence regimes
including the cooperative exchange of these technologies and
their material products, enabling their populations to grow even
more in both scale and complexity (Marlowe 2005, Hamilton et
al. 2007, Nolan and Lenski 2010, Ellis et al. 2013b).
Horticultural and agrarian societies. - The first agrarian
societies emerged in the early to mid-Holocene in more than a
dozen centers of origin spread across the continents by diverse
trajectories from coevolution in situ to cultural exchange, and
from mobile and sedentary hunter gatherers to shifting
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FIG. 4. Long-term global changes in (A) major categories of sociocultural systems (based on (Nolan and Lenski 2010), (B) human populations based on
(U.S. Census Bureau 2013), and (C) anthropogenic transformation of the terrestrial biosphere (anthrome levels; (based on Ellis et al. 2013b). Multiple
arrows indicate that Paleolithic to Neolithic transitions are regional, not global. Time scale prior to 1900 is logarithmic years BP, after 1900 is linear
calendar years.
cultivation and herding (Fuller 2010, Fuller et al. 2011, Price
and Bar-Yosef 2011, Rowley-Conwy and Layton 2011, Ellis et
al. 2013b, Fuller et al. 2014, Larson et al. 2014). The
domestication of desirable species is an especially important
form of sociocultural niche construction in which genetic
changes in populations are induced through generations of
selective breeding for desired traits in environments engineered
through land clearing and tillage, ultimately producing major
beneficial ecological inheritances for both humans and
domesticates (Smith 2007a, b, 2011, 2012, Fuller et al. 2014,
Larson et al. 2014). Through early forms of agriculture,
horticultural societies reached much greater scales of population
and cultural accumulation than that those of hunter gatherers,
and this led to further population growth, the spread of
agricultural populations into new areas and the displacement of
hunter gatherers, and to more and more complex forms of
farming and other technologies, including manuring, terracing,
and the development of smelting to produce nonferrous metal
ornaments, weapons and later other tools including farm
implements (Bellwood 2004, Bocquet-Appel 2011, Gignoux et
al. 2011). Horticultural societies developed more complex and
hierarchical forms of social organization, with some advanced
horticultural societies developing subsistence regimes based on
raiding and trade with other societies (Nolan and Lenski 2010,
Chase-Dunn and Lerro 2013).
As populations grew and agricultural and other technologies
continued to accumulate, societies continued to scale up, with
agrarian societies dependent on animal traction combined with
the plow displacing earlier horticultural and hunter gatherer
societies from the most productive lands, while also displacing
and exchanging with each other through warfare, trade,
emigration, disease, land degradation and periodic collapse
(Grigg 1974, Ellis and Wang 1997, Butzer and Endfield 2012,
Ellis et al. 2013b, Turchin et al. 2013). Over millennia, on most
continents, agrarian societies developed remarkably productive
technologies for boosting and sustaining productivity from the
same lands to support large and growing populations, including
irrigation, multiple cropping, and the use of a wide range of
fertilizers (Grigg 1974, Ellis and Wang 1997, Ellis et al. 2013b).
As populations and land productivity increased, cultural
innovations including numeracy, writing, money, the state,
absentee land ownership, and systems of unequal
intergenerational wealth transfer made possible subsistence
regimes based on the extraction of agricultural surplus by trade
and taxation, supporting the rise of non-agricultural populations
in urban settlements separated from productive lands and
specializing in craft production, trade, and the extraction of rent
from land, trade, and other subsistence resources by governing
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TABLE 6. Sociocultural subsistence regimes and their cultural, material and ecological inheritance. Benefits are italicized, detriments are bolded, and
unknown, dual consequence, and neutral in plain text. Citations add detail on subsistence strategies and may not describe inheritances.
Subsistence regime
Cultural inheritance
Cooperative hunting
social networking,
hunting
Cooking
Material
inheritance
Ecological inheritance
Citations
Human
Nonhuman
none
+foraging1 efficiency
(labor+land), overharvest
extinctions2
(Zimov et al. 1995)
fire, cooking
hearths
+nutrients from existing foods,
new foods, -disease (meat),
smoke inhalation, degraded
woodlands (fuelgathering)
+wildfire, biomass
removal
(Wrangham and ConklinBrittain 2003)
Food processing
food processing,
toolmaking
grindstones
+nutrients from existing foods,
new foods (e.g. grasses)
Hunting with lithic
projectile point
weapons
toolmaking
launchers,
projectiles
+foraging efficiency
(labor+land), overharvest
extinctions
(Shea 2006)
Landscape
modification for
hunting (corralling,
pits, wiers)
hunting, territorial
relations
stone traps
+foraging efficiency
(labor+land), steady food,
overharvest
extinctions
(Bar-Oz and Nadel 2013,
Smith 2013a)
Broadened
hunting/foraging
strategies (Broad
spectrum revolution)
biota identification, traps, storage
utilization
systems
+foraging efficiency
(labor+land), new foods,
overharvest
extinctions
(Stiner 2001, Zeder 2012)
Land clearing using
fire to enhance
hunting/foraging
burning
none
+foraging efficiency
habitat loss, extinctions, (Bliege Bird et al. 2008,
(labor+land), steady food source, invasions, habitat gain
Bowman and Haberle
2010, Rowley-Conwy
soil erosion/degradation
and Layton 2011, Smith
2011, Archibald et al.
2012, Lightfoot et al.
2013)
Propagation of
desired species
propagation
none
crops, livestock, +land
productivity, steady food
invasions
(Smith and Wishnie
2000, Smith 2007a, 2011,
Fuller et al. 2014, Larson
et al. 2014)
Artificial selection for breeding, traits
desired traits
none
crops, livestock, +land
productivity, steady food
invasions
(Smith 2007a, Fuller et
al. 2014, Larson et al.
2014)
Shifting cultivation
farming,
toolmaking
digging sticks
crops, +land productivity, steady habitat loss, invasions,
food, soil erosion/degradation
erosion, habitat gain
(Bellwood 2004, Smith
2007a)
Raiding
warfare
weapons
horses, communicable disease
invasions
(Turchin et al. 2013)
Dairy pastoralism*
livestock care
pens
livestock, steady food, pastures,
zoonotic disease, degraded
pastures
grazing competition
(Bellwood 2004)
Household livestock
(meat)
livestock care
pens, manure
storage
livestock, steady food, zoonotic
disease
habitat loss (livestock
feed)
(Larson and Fuller 2014)
Annual cultivation
farming
plows, paths
crops, weeds, soil
erosion/degradation
habitat loss
Grigg 1974)(Bellwood
2004)
Manuring
farming
manure storage
+land productivity, soil fertility,
polluted water/eutrophication
nutrient saturation
(soil, water)
Grigg 1974)
Irrigation
farming, hydrology, irrigation systems
social networking
steady food, +land productivity,
waterborne disease
salt accumulation,
water pollution,
invasions
(Grigg 1974)
Trade
tradecraft (literacy, paths, roads,
steady food, +labor productivity,
numeracy, trading), transport networks, communicable disease
social networks,
cities
prestige culture,
urban lifeways
invasions
(Smith 2004, Oka and
Kusimba 2008, Earle
2010, Feinman and
Garraty 2010, D'Odorico
et al. 2014)
Specialized crafts
craft skills,
toolmaking, trade
tools, crafts,
specialized
workplaces
steady food, +labor productivity
unknown
(Costin 1991, Costin
2001, Smith 2004)
Rent extraction from
production and/or
exchange (e.g.
governing elites,
landlords, bankers,
investors)
social organization,
infrastructure
investment, rent
extraction,
taxation
large scale
dwellings, luxury
goods, centralized
infrastructures
(ornamental,
functional)
unknown
unknown
(Smith 2004, Nolan and
Lenski 2010, ChaseDunn and Lerro 2013)
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TABLE 6. Continued.
Material
inheritance
Subsistence regime
Cultural inheritance
Industrial production
(general)
industrial sciences,
engineering,
tradecraft, social
organization
industrial
infrastructure,
industrial products
(machines to
plastics), roads,
railways
Mechanized crop
production
industrial sciences,
farming, tradecraft
Synthetic nitrogen
fertilizer
Ecological inheritance
Citations
Human
Nonhuman
steady food, surplus production,
pollution (toxic, nutrient,
carbon)
pollution (toxic,
nutrient, carbon),
extractive industrial
damage
(Basalla 1988)
Industrial
+labor productivity, energy
infrastructure,
impacts, erosion
machinery, storage
infrastructure
carbon pollution
(Grigg 1974)
nitrogen synthesis,
plant nutrition
industrial
infrastructure
+land productivity, +labor
productivity, polluted
water/eutrophication, energy
impacts
nitrogen saturation,
carbon pollution
(Smil 1991)
Pesticides
chemistry, pests
Industrial
infrastructure
steady food, +labor productivity,
resistant pests, polluted water
toxic pollution (soil,
water)
(Grigg 1974)
Service economy
service, tradecraft
communication
systems
+labor productivity
unknown
(Buera and Kaboski
2009)
*evidence of gene-culture coevolution.
1
Foraging here refers to both hunting and foraging.
2
Extinctions and invasions also include ecological inheritance from long-term population changes and trophic cascade effects on communities and
ecosystems (Estes et al. 2011).
and land-owning elites (Table 6; (Smith 2004, Nolan and
Lenski 2010, Shennan 2011b, Chase-Dunn and Lerro 2013,
Turchin et al. 2013). Technological innovations including
irrigation systems, improved roads, iron tools and weaponry,
and ships capable of sea trade further enhanced the beneficial
cultural, material and ecological inheritances of advanced
agrarian societies, aiding in their spread, their subjugation and
annexation
of other societies, and ultimately their
interconnection into a global world system by about 1600 AD
through trade, tribute and other intersocietal exchange relations
(Smith 2004, Nolan and Lenski 2010, Chase-Dunn and Lerro
2013, Turchin et al. 2013).
Urban and Industrial. - Urban populations dependent on
trade and other exchange-based subsistence regimes first arose
in the Near East more than 6000 years ago, and small cities
became common in advanced agricultural societies by 3500 BP ,
with major cities (populations >100,000) appearing by 2000 BP
(Cowgill 2004, Kirch 2005, Ellis et al. 2013b). The concentrated
populations, specialized elites, marketplaces, networks of
exchange and wealth of cities in advanced agrarian societies,
such as that of the Romans, likely facilitated remarkable cultural
and technological advances and increased incomes and
opportunities as they do in contemporary cities, despite their
relatively small share of overall population prior to the past few
centuries (Cowgill 2004, Smith 2004, Bettencourt et al. 2007,
Bettencourt and West 2010, Ortman et al. 2014, 2015). As
industrial societies arose over the past two centuries, the scale
and rate of urbanization accelerated dramatically, with the
global percentage of human populations living in cities growing
from about 7% in 1800, to 16% in 1900, to more than 50%
today (Klein Goldewijk et al. 2010). To meet the large-scale
demands of wealthy and growing urban industrial populations,
high levels of agricultural surplus production and trade were met
by ever larger scales of farming operations, trading systems, and
technological infrastructure and institutions sustained by large
energy subsidies from fossil fuels and other industrial inputs
(Grigg 1974, Lambin et al. 2001, Ellis et al. 2013b). Attendant
with the rise and centralization of large scale urban and
industrial societies has also been a continued trend toward the
systematic generation of material, cultural and political
inequality within societies and the systematic subjugation and
extraction of resources from the smaller scale and less central
societies within world systems by larger scale societies, though
these trends also include considerable rise and fall dynamics,
geographic heterogeneity and near continuous processes of
societal and ecological restructuring (Harvey 1996, Chase-Dunn
and Manning 2002, Smith 2008, Brenner and Schmid 2011,
Moore 2011, Chase-Dunn and Lerro 2013).
Evolutionary Trends in Sociocultural Niche Construction
The behaviorally modern hunter-gatherer societies of 50,000
years ago had already gained cultural and technological
capacities for ecosystem transformation beyond those of any
other multicellular species in history (Kirch 2005, Hill et al.
2009, Doughty 2013). As societies scaled up from hunter
gatherers to industrial societies, they also accumulated
technological and organizational capabilities for ecosystem
engineering and subsistence exchange that enabled their
populations to grow well beyond the capacity of unaltered
ecosystems to support them (Ellis et al. 2013b). Through the
evolution of sociocultural niche construction over hundreds of
human generations, human societies became a global force
capable of transforming the biosphere. Three main forces of
human sociocultural niche construction explain this
unprecedented capacity and its dynamics across the biosphere in
time and space; cooperative engineering, social upscaling and
energy substitution (Table 1).
The general long-term trend in sociocultural niche
construction is toward the evolution of subsistence regimes
capable of supporting ever larger and denser human populations
in increasingly unequal, hierarchical and complex societies by
increasing land productivity over time through cooperative
ecosystem engineering, increasing dependence on subsistence
exchange over larger and larger distances, and by increasing use
of nonhuman energy (Table 6). The early evolution of
increasingly productive cooperative engineering strategies and
technologies to produce more food from limited land resources
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by growing populations, or “land use intensification”, has been
explained in two ways; optimal foraging strategies and cultural
niche construction (Bird and O’Connell 2006, Price and BarYosef 2011, Shennan 2011b, Smith 2012, Ellis et al. 2013b).
Optimal foraging theory explains early forms of land use
intensification, including dietary broadening, use of fire to clear
land, and species domestication as the adoption of energy
efficient strategies for food procurement by growing populations
faced within increasingly limiting land and biotic resources,
with demographic pressures considered as the direct cause of
intensification (Bird and O’Connell 2006). Cultural niche
construction theory explains land use intensification not in terms
of demographic pressures, but “as the result of deliberate human
enhancement of resource-rich environments in situations where
evidence of resource imbalance is absent” (Smith 2012).
Contemporary archaeological evidence favors cultural niche
construction as the driver of early innovations in land use
intensification (Price and Bar-Yosef 2011, Smith 2012). In
either case, land use intensification represents the intentional
cooperative engineering of ecosystems to produce direct
beneficial ecological inheritance for human populations,
generally to the detriment of other species.
Social upscaling, centrality and urbanization
In general, societies concentrated on and used the most
productive and accessible lands first, such as lowland
floodplains, in processes of central place foraging (DysonHudson and Smith 1978, Bird and O’Connell 2006, Smith 2007a,
Price and Bar-Yosef 2011), migrating to and using more distant
and/or lower productivity lands (land use extensification) only
when population pressures built up or local resources became
degraded (Hamilton et al. 2009, Barbier 2010, Ellis et al. 2013b).
Cultural niche construction by ecosystem engineering in the
most suitable environments enabled more sedentary lifestyles,
but also induced populations to grow, requiring more productive
land-use practices to sustain them or migrations to new areas,
including wildlands (Hamilton et al. 2009, Ellis et al. 2013b).
Over time, as populations grew, societies scaled up and shifted
to subsistence regimes capable of sustaining even larger scale
societies, and these were implemented both on lands already
altered by earlier populations (central places) and through
expansion from earlier sites of settlement and land use
intensification. Subsistence regimes incapable of supporting
larger scale societies or producing detrimental cultural,
ecological or material inheritances were selected against, both
within societies, and across them by warfare, population growth
and cultural exchange.
As early societies scaled up, supported by increasingly
intensive use of the most suitable lands and by extensification of
populations across regions, the importance of central places
increased as well, as sites of denser, more resource rich, and
more culturally rich populations sustained increasingly by
cooperative strategies of subsistence exchange (Dyson-Hudson
and Smith 1978, Hamilton et al. 2009, Kaplan et al. 2009,
Burnside et al. 2012). The central places of hunter gatherers
were not cities nor were their lifestyles urbanized through
specialized subsistence regimes, the unequal distribution of
resources, hierarchical social organization, or dependence on
subsistence exchange. Yet central places still played a
significant functional role in sedentary hunter gatherer societies
as the optimal loci of networks of cultural and material exchange
(Redman 1999, Hamilton et al. 2009, Kaplan et al. 2009,
Burnside et al. 2012, Brughmans 2013, Ortman et al. 2014).
Long before the rise of cities and urban lifeways, the importance
of social networks and centrality in structuring the processes of
sociocultural niche construction, cultural and material
accumulation and subsistence exchange were established
(Dyson-Hudson and Smith 1978, Cowgill 2004, Hamilton et al.
2009, Kaplan et al. 2009, Feinman and Garraty 2010, Burnside
et al. 2012, Brughmans 2013, Ortman et al. 2014).
Over time, as societies scaled up, the populations of some
central places became the first cities, increasing in density,
wealth, inequality, social organization, cultural accumulation
and importance in focusing and guiding subsistence exchange
processes across regions (von Thünen and Schumacher-Zarchlin
1875, Christaller 1933, Stewart 1947, Redman 1999, Smith 2004,
Barbier 2010, Therborn 2011, Verburg et al. 2011, Rivers et al.
2013, Ortman et al. 2014, Smith 2014, Ortman et al. 2015). The
density of cities in itself provides advantages through the
economies of scale, increasing opportunities for wealth creation,
cultural innovation, and social connectivity, while at the same
time, increasing demands for energy to sustain this
concentration of resources and increasing potential for disease
(Redman 1999, Bettencourt 2013, Ortman et al. 2014, 2015). As
the opportunities provided by cities increase with scale, they
ultimately begin to restructure the distribution of human
populations, attracting rural immigrants in contemporary
processes of urbanization that have shifted populations from
countryside to city (Lambin et al. 2001, Klein Goldewijk et al.
2010, Smith 2014). The sustained upscaling of societies is
therefore associated with an enhancement of social centrality in
structuring and concentrating the forces of sociocultural niche
construction within landscapes and across regions and globally
through the telecoupling of resource demand and migration
(Grimm et al. 2008, Bruckner et al. 2012, Seto et al. 2012).
As societies have scaled up and increased in wealth, there has
also been a long-term trend toward increasing use of nonhuman
energy per capita; a long-term upscaling of social metabolism
(FIG 3B, Table 5)(White 1959, Smil 2008, Burnside et al. 2012,
Fischer-Kowalski et al. 2014). Beginning with the use of fire to
cook food, reducing the energy costs of digestion (Wrangham
and Conklin-Brittain 2003), humans have increasingly harnessed
nonhuman energy to support their subsistence regimes, using
harvested biomass to heat and to cook, by substituting animal
traction for human labor, and ultimately by using fossil biomass
fuels and abiotic sources of energy as substitute for labor and
even for soil fertility, through nitrogen synthesis and the mining
and transport of phosphorus and other limiting nutrients (Smil
2008, Brown et al. 2011b, Fischer-Kowalski et al. 2014). From
the beginning, energy substitution has been essential in
sustaining the upscaling of societies and their subsistence
regimes, enabling human societies to increase in scale from
small bands to telecoupled global trading systems sustained by
increasing scales of social exchange of materials, energy,
services and information.
Sociocultural niche construction as the driver of long-term
anthroecological change
Three fundamental processes of sociocultural niche
construction drive long-term ecological change. The first is
sociocultural ecosystem engineering- the ability of social groups
and societies to alter ecosystems to preferentially sustain human
populations over other species. The second is social upscaling
through culturally mediated changes in social organization and
increasing scales of subsistence exchange and third is the
harnessing of nonhuman energy sources to sustain processes of
ecosystem engineering, social upscaling and subsistence
exchange. To explore the ecological consequences of these three
processes of sociocultural niche construction as they have
unfolded across the Earth, it is critical to consider their
patterning in space and time. As societies have scaled up, so
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have rates of cultural evolution, enabling human subsistence
regimes and their transformation of ecology to evolve more
rapidly than rates of biological evolution, putting nonhuman
species at an extreme disadvantage. Social upscaling has not
only increased scales of populations and use of energy and other
resources, but has also shaped sociocultural niche construction
in space, with land use intensifying in the most suitable lands,
and populations increasingly concentrated in central places
dependent on subsistence exchange, where populations become
decoupled from direct interactions with ecosystems, while
increasing their influence at regional and global scales through
telecoupling.
ECOLOGICAL PREDICTIONS OF ANTHROECOLOGY THEORY
The most general prediction of anthroecology theory is that
the spread of behaviorally modern human societies across the
Earth caused sociocultural niche construction to emerge as a
global process of ecological change that has transformed the
terrestrial biosphere. While contemporary industrial societies
have certainly developed unprecedented capacities for
biospheric transformation (Steffen et al. 2007), evidence from
archaeology, paleoecology and environmental history confirms
that human societies have been reshaping the terrestrial
biosphere, and perhaps even global climate, for millennia (Kirch
2005, Sherratt and Wilkinson 2009, Ellis 2011, Doughty 2013,
Ellis et al. 2013b, Ruddiman 2013, Smith and Zeder 2013). Even
before the Holocene began, sociocultural niche construction by
behaviorally modern hunter-gatherers had produced major
ecological changes across Europe, Australasia and the Americas
by their hunting and trophic displacement of megafauna, causing
regime shifts in ecosystem structure (FIG 3A, (Barnosky 2008,
Estes et al. 2011, Doughty 2013, Gill 2014, Sandom et al. 2014).
The long-term upscaling, intensification, and extensification
of sociocultural niche construction would be expected to begin
as a relatively low level but widespread transformation of
ecosystems across the Earth followed by a gradually
accelerating intensification of this transformation over time, as
illustrated in FIG 3B. Contemporary evidence indicates that this
is indeed the case (Ellis et al. 2013b). Though the global extent
and dynamics of this transformation remain poorly understood,
hunter gatherer societies used fire to clear land long before the
emergence of agriculture, transforming wildlands into the open
landscape mosaics of seminatural anthromes (FIGS 3B,
4C)(Williams 2008, Ellis et al. 2013a, Ellis et al. 2013b). With
the rise of horticultural societies the first cropland anthromes
appear and then the scale, intensity and sophistication of
ecosystem engineering escalates from propagation and
domestication to the sustained tillage, irrigation, manuring and
other practices required to support the ever larger scales of
agrarian societies and later, the first urban populations subsisting
on trade and other exchange processes (FIGS 1, 3, 4)(Redman
1999, Bellwood 2004, Kirch 2005, Ellis et al. 2013b).
Though the small scale horticultural and agrarian societies of
prehistory had much lower populations than those of today, their
per capita land requirements were orders of magnitude greater
(Table 5). As a result anthropogenic transformation of the
terrestrial biosphere begins much earlier and is far more
extensive than the low populations of the past would predict
(FIGS 1, 4)(Ruddiman and Ellis 2009, Kaplan et al. 2011, Ellis
et al. 2013b). As large scale industrial societies arose, sustained
by global networks of exchange and increasing use of fossil
fuels, populations grew from 1 billion in 1800 to more than 7
billion today, ultimately transforming more than three quarters
of the terrestrial biosphere from biomes into anthromes (FIGS 1,
4)(Ellis et al. 2010, Ellis et al. 2013b). Human sociocultural
niche construction became established as a global force
transforming ecological pattern and process across the terrestrial
biosphere.
Biomes to anthromes: anthropogenic transformation
of the biosphere
A suite of more specific ecological predictions emerge from
the proposition that human sociocultural niche construction is
the main cause of long-term anthropogenic changes in
ecological pattern and process. To explore these predictions and
develop them into testable hypotheses, we begin by considering
sociocultural niche construction as a force acting on the
biosphere analogous to that of a dynamic “human climate”
interacting with ecosystems and species.
Human societies first emerged within and continue to act
upon the biomes and ecosystems formed by long-term
interactions with natural climate systems. We therefore begin by
expressing the formation of the global patterns of the biomes
and the ecosystem processes within them at regional landscape
scales (l02 to 105 km2)(Noss 1990, Ellis et al. 2012), as a
function of global variations in temperature and precipitation
acting on biota within heterogeneous terrain and soil parent
material (Pm) over time (e.g. Olson et al. 2001, Ellis and
Ramankutty 2008), as:
Biomes, ecosystems = f(temperature, precipitation,
biota, terrain, Pm, time)
(1)
To describe the formation of anthromes and
anthroecosystems through sustained processes of sociocultural
niche construction acting on the heterogeneous landscapes and
Pm of biomes, we first express the “human climate” produced
by sociocultural niche construction as function of two
sociocultural variables, society type and social centrality, and a
third variable, land suitability, as:
Sociocultural niche construction = f(society,
centrality, suitability)
with land suitability = f(society, biome, terrain)
(2)
(3)
Land suitability expresses the potential productivity of a
specific area of land in sustaining a given society, which
depends on the potential productivity attainable through
application of the societies’ ecosystem engineering practices and
subsistence regimes to a specific biome and terrain, where
terrain is considered as a factor varying within each biome,
rather than at the scale of biomes. For most societies in most
biomes, flatter areas with accessible water tend to be the most
suitable (Silbernagel et al. 1997, Huston 2005), but this can
differ greatly across societies and biomes. By combining the
variables defining sociocultural niche construction with those
defining biomes, we obtain an expression defining the formation
of anthromes and anthroecosystems over time:
Anthromes, anthroecosystems = f(biome, society,
centrality, suitability, time)
(4)
This last function provides the basis for predictions of the
anthropogenic ecological patterns and processes emerging
within a given regional landscape in response to sociocultural
niche construction by a specified society, degree of social
centrality, and land suitability. Simply put, anthroecology theory
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predicts that the ecological patterns, processes and dynamics
within and across regional landscapes inhabited by or otherwise
subject to direct interactions with behaviorally modern humans
will be predicted more accurately by equation 4 than by equation
1. In other words, anthromes will predict ecological pattern and
process more accurately than biomes alone (Ellis and
Ramankutty 2008).
Ecosystems to anthroecosystems: anthrosequences
Variations in ecological patterns and processes can be
conceptualized as “sequences”, as in chronosequences (time),
toposequences (terrain), and climosequences (climate). In this
way, “anthrosequences” (Table 1) depict variations in ecological
patterns and processes caused by variations in sociocultural
niche construction acting on a given biome. FIG 5 uses this
approach to illustrate hypothetical variations in ecological
patterns and processes in response to broad differences in types
of societies (Table 5) and variations in social centrality
(horizontal axis) and land suitability (vertical axis) across a
stylized woodland biome landscape (FIG 5A).
Patterns depicted from left to right in FIG 5 might be
interpreted as a chronosequence, and settlement patterns are
drawn to allow this. However, it must be remembered always
that societal transitions may occur in different sequences, e.g.
hunter gatherer to industrial, and that societal types are rarely
homogeneous or fully consistent. The hypothetical
anthrosequences depicted in FIG 5 are also based on patterns of
anthropogenic transformation in old world temperate woodland
biomes; very different anthrosequences would be expected in
different woodland biomes, such as tropical moist woodlands,
and in savannas, grasslands, deserts and other biomes. The
purpose here is only to demonstrate the utility of anthroecology
theory in generating testable hypotheses on anthropogenic
transformation of ecological patterns and processes based on the
three main forces of human sociocultural niche construction,
cooperative engineering, social upscaling and energy
substitution, and their spatial patterning across landscapes in
terms of social centrality and land suitability.
As illustrated in FIG 5, all types of societies alter land cover
across woodlands, fragmenting the more continuous patterns of
tree cover in natural woodlands into the complex heterogeneous
mosaics of land cover typical of most anthrome landscapes
(Ellis and Ramankutty 2008). Nevertheless, different societies
do this in very different ways, ranging from the burning of
woodland patches by more sedentary hunter gatherers, to the
wholesale clearing, cultivation, and grazing of land by agrarian
populations, to the construction of built infrastructure by dense
industrial populations and their refocusing of cultivation in the
most suitable lands remaining, abandoning the rest to woodland
recovery and grazing (FIG 5A and 5C).
The transformation of wildland biomes into anthromes by
sociocultural niche construction is depicted in FIG 5B,
illustrating a general trend towards increasingly intense use of
land for agriculture and settlements from wildlands and
seminatural lands to croplands, rangelands and dense settlements
(FIG 5C). While sociocultural niche construction is the ultimate
cause of direct anthropogenic transformation of terrestrial
ecological pattern and process, human populations and their use
of land are the proximate causes of these transformations, as
illustrated in FIG 5C. As societies increase in scale (left to right
in FIG 3 and 5; top to bottom in Table 5), human populations
become larger and denser and become more concentrated, first
into larger and larger villages and then into towns and urban
settlements of increasing size and density. Following the same
trend, extractive use of land for hunting and foraging by early
hunter gatherers transitions into increasingly intense and
complex forms of engineered ecosystem management, from the
use of fire to enhance success in hunting and foraging and to
protect the encampments of hunter gatherers, to the shifting
cultivation of crops and increasingly permanent encampments,
villages and early urban settlements of advanced horticultural
societies. Land use by agrarian and industrial societies continues
the trend towards increasingly intense and diverse use of land,
introducing continuous cropping, irrigated agriculture, and the
pasturing of livestock, and in industrial systems, the
management and conservation of forested lands, and the
introduction of ornamental land uses such as parks and yards
(FIG 5C).
Anthroecological succession and anthrobiogeography
By combining the ultimate and proximate causes of
anthropogenic transformation of terrestrial ecosystems depicted
hypothetically in FIGS 5A and 5C, relative changes in
ecosystem (FIG 5D) and biogeographic (FIG 5E) processes are
predicted in terms of sociocultural niche construction as
processes of anthroecological succession (Janzen 1983, Balée
2006, Ellis et al. 2012). Primary anthroecological succession is
the response of ecosystems and communities to their first
exposure to and transformation by sociocultural niche
construction, while secondary anthroecological succession
represents the responses of transformed ecosystems and
communities to subsequent regime shifts in societal type or
centrality. The spatial patterning and dynamics of species and
communities within and across anthromes at global and regional
scales by processes of sociocultural niche construction and
anthroecological succession is anthrobiogeography. For example,
the patterns of sociocultural niche construction depicted in FIGS
5A, 5B, 5C and their influence on habitat patch size, isolation,
megafauna biomass, plant species richness and ecosystem
novelty in FIG 5E are anthrobiogeographic patterns. The
diversity of anthropogenic ecological effects on nonhuman
species is further evident in the wide array of ecological
inheritances produced by the sociocultural subsistence regimes
listed in Table 6.
Anthroecological succession in anthromes formed by hunter
gatherer societies is caused primarily by the pressures of hunting
and foraging, together with land clearing using fire and the
propagation of desired wild plants (FIG 5). Ecological patterns
and processes strongly resemble those of wildland biomes, with
the main alterations being the formation of open woodlands and
reduced organic carbon accumulation by burning (FIG 5D), and
reductions in megafauna biomass (FIG 5E), slight increases in
species richness through the introduction of desired plant species
and enhanced rates of species introduction and establishment in
burned areas and shifts toward novel species assemblages at the
highest levels of social centrality in the most suitable lands,
where sedentary hunter gatherer settlements are located.
Sociocultural niche construction by horticultural societies
produces much more substantial transformation of ecological
pattern and process. In the least central and suitable parts of
landscapes, patterns of succession resemble those of sedentary
hunter gatherer societies. However, in the most central and
suitable parts of landscapes, shorter fallow shifting cultivation
and denser sedentary populations become established, producing
the used lands of cropland anthromes (FIG 5B). In these areas,
habitat is fragmented into smaller sized patches and
communities of exotic and domestic plants develop in response
to declining habitat isolation and size (FIG 5E) caused by
increasing social exchanges and human traffic in these areas.
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FIG. 5. Anthrosequence in a stylized temperate woodland biome illustrating conceptual relationships among society types and social centrality and their
interactions with land suitability for agriculture and settlements in shaping the spatial patterning of human populations, land use and land cover and their
ecological consequences. Settlement patterns are drawn to allow interpretation as a chronosequence of societies from left to right, however, alternate
transitions are also likely, e.g. from hunter gatherer to industrial. (A) Anthropogenic transformation of landscapes under different sociocultural systems
(top) relative to spatial variations in social centrality (horizontal axis; same for all charts below) and land suitability (vertical axis). Landscape legend is at
far left. (B) Anthrome level patterns across regional landscapes (black box frames landscape in A). (C) Variations in human population densities and
relative land use and land cover areas (white = no human use of any kind; ornamental land use = parks, yards). (D) Relative variations in ecosystem
processes, including net primary production, combustion of biomass in situ (natural fires, unintended anthropogenic fires, and intended fires, e.g., land
clearing), ex situ (hearth fires, cooking, heating), and fossil fuels, organic carbon accumulation in vegetation and soils, and reactive nitrogen and available
soil phosphorus. (E) Relative variations in biogeographic and evolutionary processes, including woodland habitat patch size and relative isolation from
other biotic communities, megafauna biomass (not including humans; native and domesticated), plant species richness of native, exotic, and domesticated
plants, and relative area of landscape without human populations or land use (wild), used directly by human populations (“used” = crops, grazing,
settlements), and transformed by human influences but not used directly (novel).
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Losses of native megafauna biomass continue, accompanied
by gains in domesticated megafauna. Biomass harvested across
landscapes for food and fuel is moved to and consumed within
settlements (ex situ combustion, hunting, gathering, crop harvest)
leading to modest accumulations of reactive N and P, enriching
soils in the vicinity of the most sedentary settlements (FIG 5D).
Agrarian societies cause far more profound shifts in
ecosystem and community patterns and processes. The larger
scales and dense populations of these societies generally
eliminate and displace native megafauna, replacing them with
domestic livestock (FIG 5E). Continuous cultivation of suitable
lands in the most socially central areas reduces primary
productivity and carbon balance through nutrient loss through
the harvest of crops for food and feed and crop residues for fuel,
causing N and P to become concentrated near settlements (FIG
5D). Low levels of agricultural productivity require near
complete cultivation of land in the most central areas where
populations are concentrated, greatly reducing habitat patch size,
almost to zero in the most central and suitable areas (FIG 5E).
This causes substantial loss of native plant species, supplanted
by smaller numbers of domesticates, and substantial exotic
establishment, mostly of weedy plants introduced through
increasing levels of social and subsistence exchange that reduce
habitat isolation (FIG 5E).
Sociocultural niche construction by industrial societies
focuses agricultural production in the most suitable landscapes
as commercial agriculture, mechanization and rural to urban
migration concentrates dense populations in cities and reduces
rural populations (FIG 5C). Use of synthetic N and mined P for
fertilizer, together with reactive N release by fossil fuel
combustion cause high levels of available N & P across
landscapes, increasing primary production across all vegetated
land cover (FIG 5D). High levels of domestic livestock are
sustained, in part through the production of feed crops. In less
suitable areas for agriculture, woodlands recover, further
enhancing net primary production. However, native plants
species are reduced significantly, and supplemented by large
numbers of exotic plant species, including high levels of
domesticates in urban areas, where large numbers of
horticultural species are maintained in yards and parks, and most
habitats are smaller in size and exposed to high volumes of
human transport, reducing their isolation from other regions to
very low levels (FIG 5E).
Nonhuman ecological inheritance and adaptations
to sociocultural niche construction
In general, the conversion of wildlands to anthromes through
sociocultural niche construction by increasingly larger scales of
societies produces both ecosystems intentionally engineered for
production and managed with increasing intensity and patches of
remnant and recovering habitats with increasing levels of
ecosystem novelty in the parts of landscapes not directly
engineered for production (FIG 5E; (Hobbs et al. 2014). To
understand the patterning of ecological communities emerging
through these patterns of anthroecological succession in
anthromes, it may be useful to consider species as comprising
functional groups in terms of their adaptations to and fitness
within the used and novel habitats of anthromes, analogous to
the grouping of species in terms of environmental adaptations
(e.g. xerophytes, mesophytes, halophytes, hydrophytes) and
guilds (e.g. frugivorous birds, insectivores, ruderals, understory
trees). For example, five simple groups might be recognized,
anthropophagics (preferred wild foods and other used wild
species not adapted to human harvest), domesticates,
anthropophiles (species tending to establish well and
outcompete other species in used and novel ecosystems),
anthropophobes (species tending not to establish in and to be
outcompeted by others in used and novel ecosystems), and
anthropoagnostics (no consistent difference in establishing
populations and competing with other species in wild vs. novel
and used ecosystems)(Bohac and Fuchs 1991, Speight and
Castella 2001, Bradley et al. 2010). By grouping species in this
way, it might be increasingly possible to predict species
assemblages in the used and novel habitats of anthrome mosaics
under different societal conditions and levels of social centrality.
Primary anthroecological succession in wildlands and
seminatural anthromes produces detrimental ecological
inheritances for anthropophagic species and their predators in all
societal types except industrial, where wild foods are uncommon.
The exceptional survival of megafauna in Africa may best be
explained by their long-term coevolution with hominins through
the gradual emergence of modern human behaviors and
sociocultural niche construction. For this reason, primary
anthroecological succession could be considered to have never
truly occurred in Africa, as species there were not confronted all
at once by behaviorally modern human societies but rather
experienced a form of secondary anthroecological succession
acting on species that may already have evolved the adaptive
traits of anthropoagnostic and/or anthropophilic species. In the
engineered used habitats of anthroecosystems, sociocultural
niche construction intentionally produces direct beneficial
ecological inheritance for domesticates, indirectly benefits
anthropophiles and filters out anthropophobes. For
anthropoagnostic species, the rise of sociocultural niche
construction has, in theory, produced no ecological inheritance
or population effects. However, as sociocultural niche
construction has increased in extent and intensity with the rise of
agrarian and industrial societies, the populations of domesticates
and anthropophiles have benefitted, while anthropophobes have
lost habitat, tending to become threatened and endangered, while
anthropophagics may be recovering, if they are not already
extinct.
Species are generally considered native based on their history
of establishment within biomes and ecoregions. Domesticates
and anthropophiles have similar relationships with anthromes
and anthroecosystems and might thus be considered native to
these in the same way. To treat such species as invading exotics
within the anthromes to which they are adapted seems both
ecologically incorrect and wildly impractical, even though this
will appear to be the case when such species become established
for the first time during early stages of primary anthroecological
succession. In considering the forces of sociocultural niche
construction as equivalent to a “human climate”, the question of
where some species “belong” shifts from ecoregions to the novel
habitats of urban landscapes, croplands, rangelands and
seminatural anthromes (e.g. (Del Tredici 2010). With the glaring
exception of most megafauna, island species, flightless birds,
and anthropophagic species, there is evidence that high levels of
both native and introduced biodiversity are generally sustained
in the multifunctional mosaic landscapes of anthromes through
high levels of biotic exchange (telecoupling; (Helmus et al.
2014), albeit in novel assemblages, habitats and ecosystems that
may have little resemblance to prior native forms (Smith and
Wishnie 2000, Kowarik 2003, Stork 2010, Kowarik 2011, Ellis
et al. 2012, Dornelas et al. 2013, Ellis 2013, Thomas 2013,
Mendenhall et al. 2014).
SHIFTING THE BASELINE: ECOLOGY IN AN
ANTHROPOGENIC BIOSPHERE
The call to integrate humans into ecology is older than the
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discipline itself and has been loud and clear for generations (e.g.
Darwin 1859, Tansley 1935, Hawley 1944, Odum 1953, Pickett
and McDonnell 1993, Redman et al. 2004, Collins et al. 2011).
Major progress has been made by theory on social-ecological
systems (Redman et al. 2004, Folke et al. 2005, Folke 2006,
Hornborg and Crumley 2007, Alessa and Chapin 2008,
Carpenter et al. 2009, Chapin III et al. 2011, Collins et al. 2011,
Levin et al. 2013), social metabolism (Baccini and Brunner 2012,
Fischer-Kowalski et al. 2014, Malhi 2014), coupled human and
natural systems (Liu et al. 2007), human ecology (Boyden 2004,
Dyball and Newell 2014), urban ecology (McIntyre et al. 2000,
Pickett et al. 2001, Grimm et al. 2008), agroecology (Tomich et
al. 2011, Gliessman 2015), countryside biogeography (Daily et
al. 2001, Mendenhall et al. 2014), novel ecosystems (Hobbs et al.
2006), anthromes (Alessa and Chapin 2008, Ellis and
Ramankutty 2008), and Anthropocene island biogeography
(Helmus et al. 2014). Anthroecology theory builds on these with
the aim of going further.
To advance the science of ecology in an increasingly
anthropogenic biosphere, it is useful to begin with “The First
Law of the Anthropocene”: the ecological patterns, processes
and dynamics of the present day, deep past, and foreseeable
future are shaped by human societies. Anthroecology theory
takes this law as given, and integrates human societies into
ecology globally across geologic time through an evolutionary
framework of human sociocultural niche construction directed at
explaining ecological pattern, process and change within and
across an increasingly anthropogenic terrestrial biosphere. In
this way, human societies are integrated into ecological science
in much the same way as climate systems, which powerfully
shape the patterns, processes and dynamics of ecology, while
also being influenced, but less so, by these interactions. As with
the classic frameworks of biogeography, succession, evolution
and ecosystem structure and function that are structured within
climate systems and the biomes formed by these, anthroecology
theory frames ecology within the “human climate” systems of
sociocultural niche construction and the anthromes formed by
these. Further, anthroecology theory suggests substantial
changes are needed in ecological science, conservation and
pedagogy if these are to advance in understanding, conserving,
and adapting to life in an increasingly anthropogenic biosphere.
Ecological science in an anthropogenic biosphere
Anthroecology theory challenges the conventional research
practices of ecological scientists in four significant ways.
The baseline is anthropogenic. - It is common practice in
ecology to define the ecological context of research sites without
incorporating either historical or contemporary social conditions.
As a result, even though most field research is conducted in sites
situated in and transformed by sustained direct interactions with
human societies, this is rarely considered or reported (Martin et
al. 2012). Given that most of the terrestrial biosphere has likely
been transformed by primary and even secondary
anthroecological succession, it is likely that the ecological
patterns and processes recognized as natural at these sites are in
fact substantially altered by sociocultural niche construction
(Rackham 1980, Cronon 1983, Redman 1999, Grayson 2001,
Briggs et al. 2006, Sih et al. 2011, Cuddington 2012, Higgs et al.
2014). While this bias might seem a minor issue, it has already
been shown to influence scientific understanding of
paleoclimate and long-term changes in community structure in
response to anthropogenic fire regimes and other forms of early
land use (Grayson 2001, Briggs et al. 2006, Jackson and Hobbs
2009, Archibald et al. 2012, Li et al. 2014). Similarly, the
tendency of ecologists to seek out the most isolated parts of
landscapes to conduct their research, such as the siting of most
permanent forest field plots in forest interiors, biases results
against understanding anthroecological pattern and process in
the small patches of trees and other fragmented habitats that now
likely represent the majority of woodlands globally (Ellis 2011).
To advance scientific understanding of ecology in an
anthropogenic biosphere, sites for field research should be
selected by more random and less biased criteria than
convenience or “the absence of readily observable
anthropogenic disturbance”. Further, the trajectory of
sociocultural conditions driving primary and secondary
anthroecological succession should be characterized and
included in the ecological descriptions of field sites in the same
way that soil type or plant association usually are, including at
minimum, the anthrome context at time of observation (see:
http://anthromes.org) and ideally, but far more challenging, the
dynamics of human populations, land use, cultural history and
social centrality starting with the earliest societies present at
sites to the present, assessed in terms of relative travel times to
nearest large settlements or markets (Verburg et al. 2011). By
including such information in ecological research, new types of
questions might be answered. For example, genomic or
metagenomic tests might determine whether there are genetic
traits associated with anthropophilia or anthropophobia, if the
anthroecological context of species is known (Allendorf et al.
2010, Sih et al. 2011).
The context is global. - Perhaps the most fundamental
challenge in understanding the anthroecological context of
observations is poor reporting of their global geographic context,
which is widespread in ecological publications (Martin et al.
2012, Dornelas et al. 2013, Karl et al. 2013). Given the free
availability of powerful geographic tools, such as Google Earth,
there is no excuse for all ecological observations not to be
described geographically, not just in the general terms of a
nearby point location, but precisely, using a polygon outlining
the extent of each observation unit for which data are reported
(Kwan 2012, Karl et al. 2013). By linking all ecological
observations with their precise geographic context, the global
and local patterns, processes, and dynamics of sociocultural
niche construction may be connected with these observations at
a later time. Further, it is useful, if possible, to make
observations across landscapes at spatial scales large enough to
represent regional to global patterns (Noss 1990) and to
investigate ecological patterns, processes and dynamics across
the full spectrum of anthromes and anthropogenic landscapes
(Martin et al. 2012). Not to do this reduces the generalizability
of ecological observations overall, and especially their relations
with anthroecological processes and their dynamics, which are
now unfolding globally through telecoupling (Kwan 2012,
Martin et al. 2012, Dornelas et al. 2013, Liu et al. 2013, Gerstner
et al. 2014, Murphy and Romanuk 2014, McGill et al. 2015).
People are data. - While the challenges of incorporating
humans into ecological research are many, so are the
opportunities. Especially exciting are recent advances in
techniques for citizen science and for sensing, cataloguing and
sharing ecological data from local to global (Fraser et al. 2013,
Crain et al. 2014, Cristescu 2014, Turner 2014, McGill et al.
2015). Macroecological study of anthroecological processes
(Burnside et al. 2012) is increasingly supported by powerful
tools for “big data” analytics, including the rise and spread of
behaviorally modern humans using paleogenomics (Pääbo 2014)
and the structure of human social networks, including changes in
social centrality (Schich et al. 2014), the crowdsourcing of
ecological questions and experiments (Fraser et al. 2013,
Sutherland et al. 2013), and new methods for integrative global
synthesis, including more powerful forms of metastudy
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(Magliocca et al. 2015) and global geospatial data and analytics
(Verburg et al. 2011, Martin et al. 2012, Schmill et al. 2014).
Ecologists have much to gain by further embracing these new
larger scale methods for socio-ecological data acquisition and
synthesis.
Ecology is a human experiment. - There are many good
reasons why it is difficult to experiment with human interactions
with ecosystems. Nevertheless, efforts to do this show
increasing promise (Felson and Pickett 2005, Felson et al. 2013)
and experimentally replicating human influences is well
developed in ecology (Debinski and Holt 2000, Fraser et al.
2013). There are real limits to these experimental approaches
however, especially in establishing cause and effect in humanenvironment interactions at larger scales and over longer
timeframes (Hedström and Ylikoski 2010, Cuddington 2012).
To develop mechanistic theory on human transformation of
ecology requires approaches that are non-deterministic,
multicausal, path-dependent and probabilistic, so as to model the
emergent anthroecological patterns and processes produced by
human individuals, groups and societies interacting with each
other in transforming ecology across landscapes and regions,
and responding to and learning from these changes (Macy and
Willer 2002, Hedström and Ylikoski 2010, Magliocca et al.
2013, Turchin et al. 2013, Magliocca et al. 2014). Agent based
modeling is ideal for this, especially using a virtual laboratory
approach applied to the anthroecological patterns observed in
real-world landscapes (Magliocca et al. 2013, 2014). These
techniques show great promise for testing basic hypotheses on
sociocultural niche construction as a force reshaping ecological
pattern and process within and across landscapes under different
societal and ecological conditions, for example the long-term
patterning of habitat quality and fragmentation under different
strategies for cooperative engineering of multifunctional
landscapes, enabling theory validation, scenario generation and
interactive assessments in the field together with stakeholders
(Matthews et al. 2007, Willemen et al. 2012).
Sustaining nonhuman nature in an anthropogenic biosphere
The challenges of sustaining nonhuman species and habitats
in an anthropogenic biosphere have never been greater as the
scale, extent, and intensity of sociocultural niche construction by
industrial societies is already without precedent and continues to
accelerate. Perhaps the greatest challenge for conserving
nonhuman species and habitats is that human harm to these is
generally not intentional, but rather results as the unintended
consequences of intentional human-benefitting sociocultural
niche construction, including ecosystem engineering for
agriculture and resource extraction (habitat loss and degradation,
pollution), industrial production and infrastructure (pollution,
hydrologic), social exchange (facilitated biotic exchange;
wildlife trade), and energy substitution (pollution, climate
change, ocean acidification). Yet the increasing global scale,
interconnection and capacity for engineering of human societies
may yet prove to be powerful forces driving major societal shifts
in both valuing and conserving nonhuman nature. The societal
benefits of sustaining nonhuman species and habitats have likely
never been clearer, as the ecological linkages among human
health, social systems and engineered environments are
increasingly understood both theoretically and with the aim of
advancing intentional management by societies (Millennium
Ecosystem Assessment 2005, Mooney et al. 2013). Perhaps the
most potent example is the recent discovery and rapid scientific
advances in understanding the novel global ecology of the
human microbiome, in which humans serve as both host
environments and biological recipients of microbial benefits and
detriments (Smillie et al. 2011, Kembel et al. 2012, O’Doherty
et al. 2014). Just as today’s globalizing and urbanizing societies
are growing more concerned with the need to conserve
nonhuman nature, they are becoming more and more capable
technologically, culturally and socially of accomplishing this
(Inglehart 2000, Rosenzweig 2003, Ellis 2015).
Engage with planetary opportunities. - The scale of human
societies is increasingly global, with telecoupling linking the
resource demands of increasingly wealthy urban populations
with transformative ecological change across the biosphere
(Lambin and Meyfroidt 2011, Fairhead et al. 2012, D'Odorico et
al. 2014, Nepstad et al. 2014). It is also highly unlikely that
human populations will decline significantly in the foreseeable
future (Bradshaw and Brook 2014). Strategies for conserving
nonhuman nature will therefore have little chance of succeeding
if they depend on halting the growth and development of human
societies. The way forward for conservation requires strategies
that can engage beneficially with global trends towards
increasing societal scales, globalization and urbanization.
Urbanization, land use intensification and decoupling are
planetary opportunities to conserve more nonhuman nature
(Fischer-Kowalski and Swilling 2011, Tilman et al. 2011,
DeFries et al. 2012). While urbanization transforms ecology
more than any other form of land use, urban areas have the
potential to be the most compact and resource efficient form of
human settlement (Grimm et al. 2008, Bettencourt and West
2010, Bettencourt 2013). As the wealth, lifestyles and other
opportunities afforded by urban living attract populations from
the countryside, opportunities are arising for woodland
recoveries in lands less suitable for industrial agriculture (Foster
et al. 1998, Rudel et al. 2009, Meyfroidt and Lambin 2011, Ellis
et al. 2013b, Queiroz et al. 2014). While the challenges are
certainly as great as the opportunities, there is good evidence
that even more land can be spared for nonhuman nature as
urbanization and societal upscaling continue, depending on the
productivity gains attainable in agriculture and forestry through
sustained land use intensification combined with more equitable
distribution to meet growing societal demands for food, feed,
housing, and energy (Neumann et al. 2010, Foley et al. 2011,
Tilman et al. 2011, Tomich et al. 2011, Loos et al. 2014). To
succeed in these efforts, it is essential to avoid the mere
displacement of societal demands from one region to another by
monitoring and governance of agricultural supply chains and
environmental programs (Fairhead et al. 2012, D'Odorico et al.
2014, Nepstad et al. 2014, Scales 2014).
Embrace change, beware domestication. - Anthropogenic
climate change, together with other indirect and direct forces of
sociocultural niche construction are causing ecological changes
that are likely more rapid than in any other recent period of
Earth history (Steffen et al. 2007, Hoffmann and Sgro 2011). To
sustain nonhuman nature, it will be necessary to assist species
and ecosystems in changing, for example by translocating
species together with their habitats as these shift towards the
poles; a very different paradigm than the classic view of
ecological conservation as the preservation of historical patterns
in situ (Bengtsson et al. 2003, Waltner-Toews et al. 2003,
Jackson and Hobbs 2009, Hobbs et al. 2011, Hoffmann and Sgro
2011, Thomas 2011, Robbins and Moore 2013, Balaguer et al.
2014, Gillson and Marchant 2014, Higgs et al. 2014, Hobbs et al.
2014, Ellis 2015). Most importantly, it will be essential to
sustain processes of evolution by natural selection in the face of
powerful human tendencies to select the traits of and even to
domesticate the native species we are trying to conserve as wild
(Western 2001, Ellis et al. 2012, Palkovacs et al. 2012, Smith et
al. 2014). The need to refocus conservation science on
sustaining evolutionary processes and their dynamics requires
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efforts and expertise that go far beyond simply preserving or
restoring historical states of ecosystems, habitats or populations,
and this need can only grow as environments become ever more
dynamic (Antrop 2006, Hoffmann and Sgro 2011, Hobbs et al.
2014), technological advances enable more precise management
of population genetics, the revival of extinct species, and the
creation of novel life forms, and these technological capacities
are confronted with expanded social demands for rewilding,
conservation beyond protected areas, and other unconventional,
controversial and poorly understood strategies for restoring and
sustaining nonhuman nature (Hobbs et al. 2011, Redford et al.
2013, Robbins and Moore 2013, Hobbs et al. 2014, Marris 2014,
Sandler 2014, Smith et al. 2014).
Bring people in: codesigning anthromes and multifunctional
landscapes. - The future of nonhuman nature depends both on
meeting human needs and inspiring human desires towards
greater efforts at Earth stewardship (Chan et al. 2007, Chapin III
et al. 2011, Marris 2011, Felson et al. 2013, Mooney et al. 2013,
Ives and Kendal 2014, Mace 2014, Palomo et al. 2014). To
accomplish this, it is more necessary than ever to integrate
sociocultural understanding into conservation (Waltner-Toews
et al. 2003, Mooney et al. 2013, Redpath et al. 2013, Ives and
Kendal 2014, Kueffer and Kaiser-Bunbury 2014, Mace 2014,
Palomo et al. 2014, Poe et al. 2014), and to consider flexible
strategies enabling the sustained integration of nonhuman
species into the novel habitats of anthromes as part of
multifunctional landscape management approaches (Antrop
2006, Bennett et al. 2006, Kleijn et al. 2011, Tomich et al. 2011,
van Noordwijk et al. 2012, Ellis 2013, Hobbs et al. 2014, Jantz
et al. 2014, Martin et al. 2014, Marvier 2014, Quinn et al. 2014).
To the extent that tradeoffs among production, biodiversity,
and ecosystem services are considered (Bergen et al. 2001,
DeFries et al. 2004, Naidoo et al. 2008) and the people and
societies with a stake in the results are involved in codesigning
and cooperating in these efforts (Berkes et al. 2000, Olsson et al.
2004, Antrop 2006, Reed 2008), multifunctional landscape
approaches have the potential to sustain nonhuman nature in the
face of unprecedented anthropogenic ecological change
(Rosenzweig 2003, DeFries and Rosenzweig 2010, DeFries et al.
2012, Ellis 2013, Hobbs et al. 2014, Martin et al. 2014).
Towards this end, ecologists will need to more actively embrace
their role in informing and helping to shape the work of
policymakers, planners, engineers and designers, as these are the
societal realms in which larger scales of human intentionality
are engaged in the processes of sociocultural niche construction
that generate ecological inheritance for nonhuman species. To
make this possible, ecologists must become more active in
observing and informing on, if not collaborating directly in, the
full range of human engagements with ecology beyond
conservation, including agriculture, industry and the built
environment, the creative and experimental processes of design
and policy, and even the decision to allow novel ecological
patterns and processes to emerge without human intervention.
Pedagogy is destiny: teach the future
With current rates of social and environmental change,
pedagogy has never been more important; teaching is the most
powerful process of social learning and is potentially capable of
shifting the cultural and ecological trajectory of societies
(Wilson et al. 2014). Moreover, the teaching of ecology and
conservation needs to change if it is to successfully assist
societies in influencing the trajectories of global
anthroecological change.
Accepting sociocultural systems as a global force of nature
represents a paradigm shift across the natural sciences that is no
less significant than evolution by natural selection or plate
tectonics (Steffen et al. 2011). For ecologists, the meaning
should be very clear. The forces of humanity are now akin to
those of climate or biology and therefore as fundamental to
understanding the processes that shape life on Earth as the
sciences of climate, soils or biology. To engage in scientific
study of ecological pattern, process and change as it exists today
and for the foreseeable future demands a firm grasp of the
human sciences and their deep integration into ecological theory
and practice. It is no longer adequate merely to study the
consequences of human transformation of ecological pattern and
process: ecology must become a science of their ultimate causes.
In the teaching of ecology, humans are generally presented as
operating entirely within a biological world, sustained by natural
ecosystems, in statements like “humanity is a biological species
in a biological world" (Wilson 2012) and “That man is, in fact,
only a member of a biotic team is shown by an ecological
interpretation of history.” (Leopold 1949). Human alteration of
ecology tends to be depicted as a recent crisis brought on by
modern industrial societies and their rapid population growth,
disturbing fragile natural ecosystems and threatening both
humanity and nonhuman nature, with such framing usually
accompanied by a call to return to or maintain some prior
balance of nature (Rockstrom et al. 2009, Simberloff 2014). As
with the seemingly perpetual need for ecology to reject the
balance of nature concept, these romantic notions should have
no place in ecological science (Cronon 1983, Briggs et al. 2006,
Pickett 2013).
There are important pedagogical consequences to this
erroneous portrayal. Beyond its incorrect interpretation of
environmental history, it implies that behaviorally modern
human populations and their transformation of ecology might be
understood, as with other species, as a matter determined simply
by population size in relation to fixed environmental limits, an
incorrect understanding of the processes that sustain societies
and cause anthropogenic ecological change (Cohen 1995,
Tainter 2006a, Ellis et al. 2013b). The ecological niche, carrying
capacity, and environmental impacts of behaviorally modern
human populations are the product of sociocultural niche
construction and are therefore defined more by sociocultural
processes than by environmental constraints (Cohen 1995,
Gurney and Lawton 1996, Odling-Smee et al. 2003c, Sayre 2008,
Ellis et al. 2013b, Odling-Smee et al. 2013). Moreover, by
singling out industrial technologies as the primary cause of
environmental harm, the fact that these are now required to
sustain existing populations is ignored, together with the fact
that these have already enabled far less land to be used per
capita over the long-term (Butzer 2012, Ellis et al. 2013b).
Further advances in ecosystem engineering efficiency combined
with more equitable subsistence regimes are the only way that
growing and thriving human populations will be able to use less
of the biosphere to produce food, fiber, energy, and other
resources. It is pedagogical malpractice to teach that
contemporary human populations might somehow sustain
themselves by going back to earlier, less efficient technologies,
such as Paleolithic lifeways, a full transition to traditional
organic farming, or an increasing dependence on harvesting
biomass for energy.
Behaviorally modern humans have always used technology to
engineer their ecosystems and have never lived in ecosystems
unaltered by their societies (Smith and Wishnie 2000, Ellis et al.
2013b). The human niche is not defined by human biology.
Humans live within a sociocultural niche constructed by
cooperative ecosystem engineering and culturally mediated
subsistence exchange. In an increasingly anthropogenic
biosphere, it is essential to shift the paradigm. Humans are a
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sociocultural species living in a sociocultural world on a used
planet. It is time to go beyond balances of nature and even
fluxes of nature to embrace the “cultures of nature” in ecology.
The paradigm must shift. Cultures create and sustain natures.
Individual humans act intentionally, but they do so within their
social contexts and depend on cultural values, perceptions, and
actions (Dyball and Newell 2014, Ives and Kendal 2014, Mace
2014, Medin and Bang 2014). The question is not how to stop
“others” from destroying nature, or finding a way to “get back to
nature”, but how to engage societies towards shaping nature
more beneficially for both humans and nonhumans (Mace 2014,
Palomo et al. 2014). Sociocultural niche construction in an
increasingly anthropogenic biosphere is neither new nor
disastrous, but the perpetual activity of human societies engaged
in the intentional cooperative engineering of ecosystems since
prehistory (Smith and Wishnie 2000, Ellis et al. 2013b).
To incorporate human sociocultural niche construction at the
core of ecological pedagogy, the framing of humans as
destroyers of nature must transition to narratives of societies as
nature sustainers (Chapin III et al. 2011). In moving towards this
goal, the work of archaeologists, natural historians,
agroecologists, urban ecologists, conservationists, engineers and
designers all have much to offer and much to gain.
The teaching of ecology has always appealed to a sense of
wonder about the natural world. As educators we must build a
new sense of wonder and discovery about the “tangled bank” of
human sociocultural systems and the diversity of ecosystems
they create and sustain together with the traditional ecologies of
cultures, cultural landscapes and cities. By embracing
sociocultural evolution and teaching it, current and future
generations of ecologists and the public will be better equipped
to guide societies towards better outcomes for both people and
nonhuman nature.
Thinking globally in an anthropogenic biosphere
The call to recognize the Anthropocene as a new epoch of
geologic time confronts ecologists and other environmental
scientists with the need to understand, visualize and model the
emergence and dynamics of human societies as a global force
reshaping the biosphere, atmosphere and the other “spheres” of
the Earth system (Ellis 2011, Steffen et al. 2011). Anthroecology
characterizes these global forcings and dynamics through the
analogy of a “human climate system”. Just as Earth’s climate
system shapes the dynamics of energy and material flow across
the atmosphere, hydrosphere and other spheres, human systems
shape the dynamics of energy, material, biotic and information
flow across the biosphere and other spheres, including those of a
newly emerged anthroposphere comprised of human societies
and their material cultures (Ellis and Haff 2009, Lucht 2010,
Steffen et al. 2011, Baccini and Brunner 2012). Human systems
and their interactions with the biosphere and anthroposphere are
responsive to feedbacks with other Earth systems and are
dynamic in response to evolutionary changes in sociocultural
niche construction. In this way, long-term changes in human
social organization, cooperative ecosystem engineering,
exchange relationships, and energy systems are coupled with
long-term changes in the Earth system. It remains to be seen
whether intentional efforts by societies to intervene in the
dynamics of human systems at global scales can or will
ultimately generate more beneficial and less detrimental
ecological inheritance for both human societies and nonhuman
species.
CONCLUSIONS: THE NATURES WE CREATE
Behaviorally modern humans are Earth’s first ultrasocial
species, requiring social learning to survive and to reproduce
within the sociocultural systems and cooperatively engineered
ecosystems that sustain them. Behaviorally modern human
societies began transforming terrestrial ecology more than
50,000 years ago and emerged as a global force as their
populations spread out of Africa and across the Earth. While
contemporary rates and scales of anthropogenic ecological
change are unprecedented, behaviorally modern human societies
began permanently reshaping the terrestrial biosphere countless
generations before the rise of industrial societies.
This paper introduces a causal theory explaining the
emergence and dynamics of human transformation of the
biosphere based on sociocultural niche construction, an
evolutionary theory combining socially learned cooperative
ecosystem engineering, the upscaling of societies through
culturally mediated changes in social organization and
subsistence exchange, and the harnessing of nonhuman energy
sources to sustain these processes. In developing this theory,
archaeological, paleoecological, anthropological, sociological,
historical and evolutionary evidence have been presented
demonstrating that the ultimate causes of human transformation
of the biosphere are inherently social and cultural, not biological,
chemical or physical.
The emergence of sociocultural niche construction by
behaviorally modern human societies represents a novel
evolutionary process in the Earth system that has reshaped the
biosphere and will likely continue reshaping both the biosphere
and human societies for the foreseeable future. Building on this
"first law of the Anthropocene”, anthroecology theory generates
novel ecological hypotheses together with strategies for testing
them using new theoretical frameworks including
anthroecosystems, anthrosequences, anthrobiogeography and
anthroecological succession. By engaging these frameworks
together with sociocultural niche construction theory, the
emergence and long-term dynamics of anthropogenic ecological
patterns and processes in biogeography, ecological succession,
ecosystems and landscapes, including the reshaping of biomes
into anthromes, can be more effectively investigated and
understood in an increasingly anthropogenic biosphere.
Ultimately, anthroecology theory aims to shift the science
and pedagogy of ecology beyond the classic paradigm of
“natural systems with humans disturbing them” to a new
paradigm of “societies sustaining an anthropogenic biosphere”.
In applying anthroecology theory it is critical to remember that
like biological evolution, sociocultural evolution is a process,
not a destiny, and that the future remains fully open to surprise.
Perhaps the only guarantee is that the future will likely include
societies and ecosystems that bear little resemblance to those of
today. Nevertheless, it is hoped that as ecological science
advances in its capacity to investigate and understand the
ultimate causes, not just the consequences, of human
transformation of the biosphere, that this capacity will help to
guide societies towards sustaining nonhuman natures more
successfully in a thriving anthropogenic biosphere that future
generations across the world will be proud of.
ACKNOWLEDGEMENTS
The author thanks Aaron Ellison for inviting this work with
the question: how would ecological concepts and ideas have to
change if we (re)focused our attention on anthromes, not biomes,
as an underlying biogeographic organizing schema? This work
would not have been possible without the sustained support of
Ariane de Bremond. The author also thanks Jared Margulies,
Nicholas Magliocca, David Lansing, Robert Dyball, Michael
Perring, and three anonymous reviewers for their very helpful
Page 27 of 40
Ellis, E.C. 2015. Ecology in an Anthropogenic Biosphere. Ecological Monographs in press doi.org: 10.1890/14-2274.1 [FORMATTED PREPRINT]
comments on the manuscript. Support for Erle Ellis was
provided by US NSF grants CNS 1125210 and DBI 1147089.
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